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SANTA   BARBARA.   CALIFORN 


GRAY'S  BOTANICAL  TEXT-BOOK 

VOLUME  I. 
STRUCTURAL    BOTANY 


GRAY'S  BOTANICAL  TEXT-BOOK 

CONSISTS  or 

VOL.  I.    STRUCTURAL  BOTANY.    By  ASA  GRAY. 

II.    PHYSIOLOGICAL  BOTANY.    By  GEORGE  L.  GOODALE. 

III.  INTRODUCTION    TO     CRYPTOGAMIC    BOTANY,     BOTH 

STRUCTURAL  AND  SYSTEMATIC.     By  WILLIAM  G. 
FARLOW.     (In  preparation.) 

IV.  SKETCH  OF  THE  NATURAL  ORDERS  OF  PH^NOGAMOUS 

PLANTS;  their  Special  Morphology,  Useful  Pro- 
ducts, &c.     (In  preparation.) 


O-RAY'S     BOTANICAL     TEXT-BOOK. 

(SIXTH  EDITION.) 

VOL.  I. 


STRUCTURAL    BOTANY, 

OK 

ORGANOGRAPHY  ON  THE  BASIS  OF 

MORPHOLOGY. 


TO  WHICH  IS  ADDED  THB  PRINCIPLES  OF 


TAXONOMY  AND  PHYTOGRAPHY, 


of  -Botanical  Cermet. 


BY 

ASA  GRAY,  LL.D.,  ETC., 

FISHER  PROFESSOR  OP  NATURAL  HISTORY  (BOTANY)  III 
HARVARD  UNIVERSITY. 


NEW  YORK    •:•    CINCINNATI    •:•    CHICAGO 

AMERICAN     BOOK     COMPANY 


GRAY'S    BOTANICAL   SERIES 


Gray's  How  Plants  Grow 

Gray's  How  Plants  Behave 

Gray's  Lessons  in  Botany 

Gray's  Field,  Forest,  and  Garden  Botany 

(Flora  only) 

Gray's  School  and  Field  Book  of  Botany 

(Lessons  and  Flora) 

Gray's  Manual  of  Botany.    (Flora  only) 
Gray's  Lessons  and  Manual  of  Botany 

Gray's  Botanical  Text-Book 
I.     Gray's  Structural  Botany 
II.     Goodale's  Physiological  Botany 

Coulter's  Manual  of  Botany  of  the  Rocky 

Mountains 
Gray   and    Coulter's   Text-Book    of 

Western  Botany 


Copyright,  1879,  by  ASA  GRAY 
Copyright,  1907,  by 

PRESIDENT  AND  FELLOWS  OF  HARVARD  COLLEGE 

OKAY'S  STRUCTURAL  BOTANY 
w.  p.  ;i3 


MS 


PREFACE. 


THE  first  edition  of  this  treatise  was  published  in  the 
year  1842,  the  fifth  in  1857.  Each  edition  has  been  in 
good  part  rewritten, — the  present  one  entirely  so, —  and  the 
compass  of  the  work  is  now  extended.  More  elementary 
works  than  this,  such  as  the  author's  First  Lessons  in 
Botany  (which  contains  all  that  is  necessary  to  the  prac- 
tical study  of  systematic  Phsenogamous  Botany  by  means 
of  Manuals  and  local  Floras),  are  best  adapted  to  the 
needs  of  the  young  beginner,  and  of  those  who  do  not 
intend  to  study  Botany  comprehensively  and  thoroughly. 
The  present  treatise  is  intended  to  serve  as  a  text-book  for 
the  higher  and  completer  instruction.  To  secure  the 
requisite  fulness  of  treatment  of  the  whole  range  of  sub- 
jects, it  has  been  decided  to  divide  the  work  into  distinct 
volumes,  each  a  treatise  by  itself,  which  may  be  indepen- 
dently used,  while  the  whole  will  compose  a  comprehensive 
botanical  course.  This  volume,  on  the  Structural  and 
Morphological  Botany  of  Phsenogamous  Plants,  properly 
comes  first.  It  should  thoroughly  equip  a  botanist  for  the 
scientific  prosecution  of  Systematic  Botany,  and  furnish 
needful  preparation  to  those  who  proceed  to  the  study  of 
Vegetable  Physiology  and  Anatomy,  and  to  the  wide  and 
varied  department  of  Cryptogamic  Botany. 


iv  PREFACE. 

The  preparation  of  the  volume  upon  Physiological 
Botany  (Vegetable  Histology  and  Physiology)  is  assigned 
to  the  author's  colleague,  Professor  GOODALE. 

The  Introduction  to  Cryptogamous  Botany,  both  structu- 
ral and  systematic,  is  assigned  to  his  colleague,  Professor 
FARLOW. 

A  fourth  volume,  a  sketch  of  the  Natural  Orders  of 
Phaenogamous  Plants,  and  of  their  special  Morphology, 
Classification,  Distribution,  Products,  &c.,  will  be  needed 
to  complete  the  series:  this  the  present  author  may 
rather  hope  than  expect  himself  to  draw  up. 

ASA  GRAY. 

HERBARIUM  OF  HARVARD  UNIVERSITY, 
CAMBRIDGE   April  10, 1879. 


%*  The  numerals  in  parentheses,  which  are  here  and  there  introduced 
into  sentences  or  appended  to  them,  are  references  to  the  numbered  para- 
graphs in  which  the  topic  is  treated  or  the  term  explained. 


CONTENTS. 


PAGK 

INTRODUCTION.    THE  DEPABTMENTS  OF  THE  SCIENCE    ....       1 


CHAPTER  I.  OUTLINES  OF  THE  GENERAL  MORPHOLOGY 

OF  PH^ENOGAMOUS  PLANTS 6 

CHAPTER  II.  MORPHOLOGY  AND  DEVELOPMENT  OF  THE 

EMBRYO  AND  SEEDLING 9 

The  Embryo,  its  Nature,  Structure,  and  Parts • 

Development  of  the  Dicotyledonous  Embryo  in  Maple   ...  10 

In  Ipomoea,  or  Morning  Glory,  &c.,  with  Albuminous  Seeds    .  13 

In  Embryos  with  thickened  Cotyledons If 

As  of  Almond,  Beech,  Bean,  &c 17 

With  Hypogseous  Germination  and  no  Elongation  of  Caulicle  19 

In  Megarrhiza,  &c.,  with  concreted  Petioles  to  the  Cotyledons  21 
In  Ipomoea    leptophylla   with   foliaceous    and    long-petioled 

Cotyledons  and  no  elongation  of  Caulicle 22 

In  Pumpkin,  &c.,  with  no  Primary  Root 22 

The  Polycotyledonous  Embryo 23 

The  Monocotyledonous  Embryo  of  Iris,  Onion,  Cereal  Grains  24 

Pseudo-monocotyledonous  and  Acotyledonous  Embryo  ...  26 

Dicotyledonous  and  Monocotyledonous  Plants 27 

CHAPTER  III.  MORPHOLOGY  AND  STRUCTURE  OF  THE 

ORGANS  OF  THE  PLANT  IN  VEGETATION   .  .  27 

SECTION  I.    OF  THE  ROOT 27 

Nature,  Growth,  and  Composition 28 

Root-hairs 29 

Kinds  of  Roots 29 

Duration;  Annuals 30 

Biennials 31 

Perennials 32 

Aerial  Roots       33 

Epiphytes  or  Air-plants 35 

Parasitic  Plants,  Green  and  Colored 36 


fl  CONTENTS. 

SBCTIOX  n.    Or  BUDS 40 

Scaly  Buds  and  Bud-scales 40 

Naked,  Subpetiolar,  and  Fleshy  Buds 41 

Bud-propagation 43 

Normal,  Accessory,  and  Adventitious  Buds 44 

SECTION  III.    OF  THE  STEM 46 

§  1.    GENERAL  CHARACTERISTICS  AND  GROWTH 45 

Development  and  Structure 46 

Ramification,  Branches 47 

Excurrent  and  Deliquescent  Stems 48 

Definite  and  Indefinite  Annual  Growth 49 

§  2.    FORMS  OF  STEM  AND  BRANCHES 50 

Herbs,  Shrubs,  Trees,  Culm,  Caudex,  Scape 50 

Climbing  Stems,  Twining  or  otherwise 61 

Leaf-climbers,  Tendril-climbers,  and  Root-climbers     ....  62 

Suckers,  Stolons,  Offsets,  Runners 53 

Tendrils  formed  of  Stems 64 

Sympodial  and  Monopodial  Stems 66 

Spines  or  Thorns  and  Subterranean  Stems 56 

Rhizoma  or  Rootstock 67 

Tuber,  Tubercles 69 

Corm  or  Solid  Bulb 61 

Bulb,  Bulblets 62 

Condensed  Aerial  Stems 64 

Stems  serving  for  Foliage,  Phyllocladia,  Cladophylla     ...  66 

Frondose  Stems 66 

§  3.    INTERNAL  STRUCTURE 67 

Anatomical  Elements 68 

Endogenous  Structure 70 

Exogenous  Structure ;  its  Beginning 73 

First  Year's  Growth 74 

Pith,  Layer  of  Wood,  &c 75 

Bark,  its  Parts  and  Structure       76 

Annual  Increase  in  Diameter 78 

Demarcation  of  Annual  Layers 79 

Sap-wood  and  Heart-wood 80 

Growth  and  Duration  of  Bark 81 

Living  Parts  of  a  Tree  or  Shrub,  Longevity 83 

The  Plant  composite 84 

SECTION  IV.    OF  LEAVES 86 

§  1.    THEIR  NATURE  AND  OFFICE 85 

Parts  of  a  Leaf 85 

Duration,  Defoliation,  Normal  Position 86 


CONTENTS.  Til 

§  2.    THEIR  STRUCTURE  AND  FORMS  AS  FOLIAGE 87 

Internal  Structure  or  Anatomy 87 

Parenchyma-cells 88 

Epidermis,  Stomata  or  Breathing-pores 89 

Framework,  Venation 90 

Parallel-veined  or  Nerved  Leaves 91 

Keticulated  or  Netted-veined  Leaves 92 

Pinnately  or   Feather-veined   and    Palmately  or  Radiately 

Veined 93 

Forms  as  to  Outline 94 

Forms  as  to  Extremity 96 

Forms  as  to  Margin  or  Special  Outline  and  Dentation  ...  97 

Lobation  or  Segmentation 98 

Number  and  Arrangement  of  Parts 99 

Compound  Leaves,  Pinnate  and  Palmate  or  Digitate,  &c.    .    .  100 

Petiole  or  Leafstalk     .„ 104 

Stipules,  Ligule,  Stipels 106 

Leaves  in  unusual  Modifications 106 

Such  as  Inequilateral,  Connate,  Perfoliate 107 

Vertical  and  Equitant 108 

Without  distinction  of  Parts 109 

Stipules  serving  for  Blade 109 

Phyllodia,  or  Petioles  serving  for  Blade 110 

§  3.    LEAVES  SERVING  SPECIAL  OFFICES 11CP 

Utilizing  Animal  Matter 110 

Ascidia  or  Pitchers Ill 

Sensitive  Fly-traps 113 

Leaves  for  Storage 116 

Bulb-scales  and  Bud-scales 116 


CHAPTER  IV.    PHYLLOTAXY,  OR  LEAF-ARRANGEMENT  .  119 

SECTION  I.    DISTRIBUTION  OF  LEAVES  ON  THE  STEM   ....  119 

Phyllotaxy  either  Verticillate  or  Alternate,  Cyclical  or  Spiral  119 

Verticillate  or  Cyclical  Arrangement 120 

Alternate  or  Spiral  Arrangement 121 

Its  Modes  and  Laws 122 

Relation  of  Whorls  to  Spirals 129 

Hypothesis  of  the  Origin  of  Both 130 

Fascicled  Leaves 131 


SECTION  II.    DISPOSITION  OF  LEAVES  IN  THE  BUD 


Vernation  and  ^Estivation ;  the  Modes 132 

Direction,  Dextrorse  and  Sinistrorse 140 


Vili  CONTENTS. 

CHAPTER  V.    ANTHOTAXY  OR  INFLORESCENCE   ....  141 

Bracts  and  Bractlets  and  their  Modifications 141 

Peduncles,  Pedicels,  Rhachis,  Receptacle 143 

Position  of  Flower-buds,  Kinds  of  Inflorescence 144 

Indeterminate,  Indefinite,  or  Botryose 146 

Raceme,  Corymb,  Umbel 146 

Head  or  Capitulum 147 

Syconium  or  Hypanthodium 148 

Spike,  Spadix,  Arnent  or  Catkin 149 

Panicle  and  other  Compound  Forms 160 

Determinate  or  Cymose 151 

Cyme,  Glomerule,  &c       152 

Botryoidal  Forms  of  Cymose  Type 153 

Sympodial  Forms 154 

Scorpioid  and    Helicoid,  the  Pleiochasium,  Dichasium,  and 

Monochasium 156 

Bostryx,  Cincinnus,  Rhipidium,  Drepanium,  &c 166 

Mixed  Inflorescence 168 

Thyrsus,  Verticillaster,  &c 169 

Relations  of  Bract,  Bractlet,  and  Flower 160 

Anterior  and  Posterior,  or  Inferior  and  Superior 160 

Median  and  Transverse 160 

Position  of  Bractlets 161 

Tabular  View  of  Inflorescence 162 

CHAPTER  VL    THE  FLOWER 163 

SECTION  I.    ITS  NATURE,  PARTS,  AND  METAMORPHT     ....  163 

Floral  Envelopes,  Perianth,  or  Perigone 164 

The  Parts,  Calyx  and  Corolla 165 

Androecium,  Stamens 166 

Gynoecium,  Pistils 166 

Torus  or  Receptacle  of  the  Flower 167 

Metamorphosis 167 

Unity  of  Type  illustrated  by  Position  and  Transitions    ...  169 

Teratological  Transitions  and  Changes 170 

SECTION  II.    FLORAL  SYMMETRY 174 

Symmetrical,  Regular,  and  Complete  Flower 175 

Numerical  Ground-plan 176 

Pattern  Flowers 176 

Diplostemonous  Type 177 

SECTION  III.    VARIOUS  MODIFICATIONS  OF  THE  FLOWER  .    .    .  179 

§  1.    ENUMERATION  OF  THE  KINDS 179 

J2.    REGULAR  UNION  OF  SIMILAR  PARTS 180 

Coalescence  or  Cohesion       180 


CONTENTS.  iX 

§  3.    UNION  or  DISSIMILAR  OR  SUCCESSIVE  PARTS    ....  181 

Adnation  or  Connation 182 

Hypogynous,  Perigynous,  Epigynous 183 

5  4.    IRREGULARITY  OF  SIMILAR  PARTS 184 

§  6.    DISAPPEARANCE  OR  OBLITERATION  OF  PARTS     ....  187 

Abortion  or  Suppression  of  Parts  of  a  Circle 187 

Abortion  or  Suppression  of  whole  Circles 190 

Terms  therewith  connected 191 

Suppressed  Perianth 191 

Suppressed  Andrcecium  or  Gynoecium 193 

Along  with  suppressed  Perianth 194 

Neutral  Flowers 195 

§  6.    INTERRUPTION  OF  NORMAL  ALTERNATION 196 

Anteposition  or  Superposition 196 

In  Appearance  only 196 

Superposition  by  Spirals 198 

Anteposition  with  Isostemony  and  Diplostemony 197 

With  Obdiplostemony 198 

§  7.    INCREASED  NUMBER  OF  PARTS 200 

Regular  Multiplication 200 

Parapetalous  Multiplication 201 

Chorisis  or  Deduplication 202 

§  8.    OUTGROWTHS 209 

Their  relation  to  Chorisis :  Trichomes 209 

Corona  or  Crown 210 

Ligule 211 

§  9.    FORMS  OF  THE  TORUS  OR  RECEPTACLE 211 

Stipe,  Thecaphore,  Gynophore,  Carpophore,  &c. 212 

Disk 213 

Hypanthium 214 

SECTION  IV.     ADAPTATIONS  OF  THE  FLOWER  TO  THE  ACT  OF 

FERTILIZATION 216 

§  1.    IN  GENERAL 215 

Close  and  Cross  Fertilization,  or  Autogamy  and  Allogamy     .  216 

§  2.    ADAPTATIONS  FOR  ALLOGAMY  OR  INTERCROSSING   .    .    .  216 

Wind-fertilizable  or  Anemophilous  Flowers 217 

Insect-fertilizable  or  Entomophilous  Flowers 218 

Irregularity  as  related  to  Allogamy 219 


CONTENTS. 

Dichogamy,  either  Proterandrous  or  Proterogynou»  ....  219 

Proterogyny 219 

Proterandry 220 

Particular  Adaptations  in  Papilionaceous  Flowers      ....  225 

In  Kalmia-blossoms,  Iris,  &c 229 

Transportation  of  Pollinia 230 

In  Orchidaceae  and  Asclepiadaceae 231 

Heterogonous  Dimorphism  and  Trimorphism 234 

§  3.    ADAPTATIONS  FOR  CLOSE  FERTILIZATION 240 

Cleistogamy 241 

SECTION  V.    THE  PERIANTH,  OR  THE  CALYX  AND  COROLLA  IN 

PARTICULAR 243 

Perianth  as  to  Duration,  Numerical  Terms,  Union,  &c.  .     .    .243 

Parts  of  Petals  and  of  Gamophyllous  Perianth 246 

Forms  of  Corolla  and  Calyx 246 

SECTION  VI.    THE  ANDROSCIUM,  OR  STAMENS  IN  PARTICULAR  .  249 

The  Stamen  as  a  whole  ;  Numerical  Terms        249 

The  Filament  and  the  Anther;  their  Modifications    ....  251 

Pollen 266 

Pollen-tubes 268 

SBCTION  VII.    THE  PISTILS,  OR  GTN<ECIUM 269 

§  1.    IN  ANGIOSPERMS 269 

Carpel  or  Carpophyll 260 

Ventral  and  Dorsal  Sutures ;  Placenta 261 

Simple  or  Apocarpous  Pistils 262 

Compound  or  Syncarpous  Pistil 263 

With  two  or  more  Cells  and  Axile  Placentae ;  Partitions    .    .  264 

With  one  Cell  and  Parietal  Placentae 265 

With  one  Cell  and  Free  Central  Placenta 266 

Anomalous  Placentation 267 


§  2.    IN  GTMNOSPERMS 268 

Structure  in  Gnetaceae 269 

Structure  in  Coniferae 270 

In  the  Yew  Family 271 

In  the  Pine  Tribe,  &c 272 

In  the  Cypress  Tribe 273 

Structure  in  Cycadaceae 274 

SECTION  VTIL    THE  OVULE 276 

Its  Structure  and  Position 277 

Its   Forms,    Orthotropous,  Campylotropou*,    Amphitropous, 

Anatropous 278 

Origin  and  Morphological  Nature  of  the  Ovule 282 

Origination  of  the  Embryo 283 


CONTENTS.  » 

CHAPTER  VII    THE  FRUIT 286 

SECTION  I.     ITS  STRUCTURE,  TRANSFORMATIONS,  AND   DBHIS- 

CENCE 286 

Pericarp,  its  Alterations,  Accessions,  and  Transformations     .  287 

Dehiscence 288 

SECTION  II.    THE  KINDS  OF  FRUIT 291 

Simple  Fruits 291 

Dehiscent  Fruits,  Follicle,  Legume,  Capsule,  Pyxis,  Silique    .  292 
Indehiscent  Dry  Fruits,  Samara,  Akene,  Utricle,  Caryopsis, 

Nut,&c 294 

Fleshy  Fruits,  Drupe,  Pome,  Pepo,  Berry,  &c 297 

Aggregate  Fruits 299 

Accessory  or  Anthocarpous  Fruits 300 

Multiple  or  Collective  Fruits,  Syconium,  Strobile,  &c.    ...  301 

Table  of  Simple  Fruits 304 

CHAPTER  Vin.    THE   SEED 305 

Its  Stalk,  Coats,  and  Appendages 306 

Aril  or  Arillus 308 

Nucleus  or  Kernel,  Albumen 309 

The  Embryo,  its  Parts  and  Positions 311 

The  Cotyledons  as  to  Adjustment  and  Number 313 

CHAPTER  IX.    TAXONOMY 315 

SECTION  I.    THE  PRINCIPLES  OF  CLASSIFICATION  IN  NATURAL 

HISTORY 316 

Individuals 316 

Species 817 

Varieties,  Races,  &c 318 

Cross-breeds  and  Hybrids 321 

Genera 323 

Orders,  Classes,  Tribes,  &c 326 

Sequence  of  the  Grades 327 

Nature  and  Meaning  of  Affinity 327 

Theory  of  Descent  and  Natural  Selection 328 


SECTION  II.    BOTANICAL  CLASSIFICATION 


Ante-Linnaean  Classifications 332 

Linnaean  Classification 333 

Sexual  Artificial  System 334 

Natural  System J38 

As  presented  by  Jussieu 339 

Some  of  its  Modifications 340 


Xii  CONTENTS. 

CHAPTER  X.    PHYTOGRAPHY 846 

SECTION  I.    NOMENCLATURE 846 

Names  of  Plants,  Binomial  Nomenclature 346 

Rules  for  naming  Plants 847 

Names  of  Genera 348 

Names  of  Species,  Varieties,  &c 350 

The  Fixation,  Precision,  and  Citation  of  Names  ' 362 

Subgeneric  Names 356 

Tribal  and  Ordinal  Names 367 

Names  of  Cohorts,  Classes,  &c 358 


SECTION  II.    GLOSSOLOGY  OR  TERMINOLOGY 


SECTION  III.    DESCRIPTION  ..............  861 

Characters     ..................  361 

Punctuation  ..................  364 

Synonomy      ..................  365 

Iconography  ..................  366 

Habitat  and  Station,  &c  ..............  366 

Etymology  of  Names  ...............  366 

Accentuation,  Abbreviations    ............  367 

Signs     ....................  368 

Floras,  Monographs,  &c  ..............  369 

SECTION  IV.     SPECIMENS,  DIRECTIONS  FOR  THEIR   EXAMINA- 

TION, PRESERVATION,  &c  .............  870 

Implements  of  Investigation    ............  370 

Diagrams  ...................  371 

Herborizing    ..................  371 

Drying  Specimens  ................  375 

Poisoning  Specimens  ...............  379 

The  Herbarium  .................  380 


ABBREVIATIONS     ................    386 

SIGNS    ...........  .  ..........    891 

GLOSSARY  OF  BOTANICAL  TERMS,  WITH  INDEX     .    .    393 


STRUCTURAL    BOTANY 


THE    BASIS    OF    MORPHOLOGY. 


INTRODUCTION. 

1.  THE  two  Biological  Sciences,1  considered  as  parts  of  Natural 
History,  are  Zoology  and  Botany.     The  latter  is  the  natural 
history  of  the  Vegetable  Kingdom.     It  embraces  every  scientific 
inquiry  that  can  be  made  respecting  plants,  their  nature,  their 
kinds,  the  laws  which  govern  them,  and  the  part  they  play  in 
the  general  economy  of  the  world. 

2.  We  cannot  distinguish  the  vegetable  from  the  animal  king- 
dom by  any  complete  and  precise  definition.     Although  ordinary 
observation  of  their  usual  representatives  may  discern  little  that 
is  common  to  the  two,  yet  there  are  many  simple  forms  of  life 
which  hardly  rise  high  enough  in  the  scale  of  being  to  rank  dis- 
tinctively either  as  plant  or  animal ;  there  are  undoubted  plants 
possessing  faculties  which  are  generally  deemed  characteristic  of 
animals ;  and  some  plants  of  the  highest  grade  share  in  these 
endowments.     But  in  general  there  is  a  marked  contrast  between 
animal  and  vegetable  life,  and  in  the  part  which  animals  and 
plants  respectively  play  in  nature. 

3.  Plants  only  are  nourished  upon  mineral  matter,  upon  earth 
and  air.     It  is  their  peculiar  office  to  appropriate  mineral  mate- 
rials and  to  organize  them  into  a  structure  in  which  life  is  mani- 
fested, —  into  a  structure  which  is  therefore  called  organic.     So 
the  material  fitted  for  such  structure,  and  of  which  the  bodies 

1  Biology,  the  science  of  life,  or  rather  of  living  things,  in  its  earlier  use 
was  equivalent  to  physiology :  recently,  it  has  come  to  denote  the  natural 
history  of  plants  and  animals,  f.  e.  of  the  two  organic  kingdoms,  including 
both  their  physiology  and  descriptive  natural  history, 

1 


2  LNTKODTJCTION. 

of  plants  and  animals  are  composed,  is  called  organic  matter. 
Animals  appropriate  and  live  upon  this,  but  have  not  the  power 
of  producing  it.  So  the  vegetable  kingdom  stands  between  the 
mineral  and  the  animal ;  and  its  function  is  to  convert  materials 
of  the  one  into  food  for  the  other.  Although  plants  alone  are 
capable  of  building  up  living  structure  out  of  mineral  mate- 
rials, and  are  the  sole  producers  of  the  organic  matter  which 
is  essential  to  animal  life,  and  although  animals  consume  that 
which  plants  produce,  yet  plants  also  consume  organic  matter, 
more  or  less,  acting  in  this  respect  like  animals  in  all  their  opera- 
tions, except  in  the  grand  and  peculiar  one  by  which  they 
assimilate  mineral  matter.  Most  plants  of  the  higher  grades 
assimilate  largely  and  consume  little,  except  in  special  opera- 
tions. Some,  on  the  contrary,  are  mainly  consumers,  and  feed 
upon  formed  organic  matter,  living  in  this  respect  after  the 
manner  of  animals.  The  living  substance  of  plants  and  animals 
is  essentially  the  same. 

4.  Botany  deals  with  plants  :  1.  As  individuals,  and  in  respect 
to  their  structure  and  functions.     2.  In  their  kinds,  and  as 
respects  their  classification,  nomenclature,   &c.      Accordingly, 
the  most  comprehensive  division  of  the  science  is  into  PHYSIO- 
LOGICAL or  BIOLOGICAL  BOTANY  (using  these  terms  in  their  widest 
sense)  and  SYSTEMATIC  BOTANY.    But  as  Physiology  and  Biology, 
in  the  restricted  sense,  relate  only  to  functions  or  actions  and 
their  consequences,  the  first  department  naturally  divides  into 
two,  viz.  Structural  Botany  and  Physiology. 

5.  STRUCTURAL  BOTANY  comprehends   all  inquiries  into  the 
structure,  the  parts,  and  the  organic  composition  of  vegetables. 
This  is  termed  ORGANOGRAPHY,  when  it  considers  the  organs  or 
obvious  parts  of  which  plants  are  made  up,  and  MORPHOLOGY, 
when   the   study  proceeds   on   the   idea   of  type.      The  term 
ORGANOGENY   has   been   applied  to   the   study  of  the  nascent 
organs    and    their    development ;    PHYTOTOMY,    or  VEGETABLE 
ANATOMY,  to  that  of  the  minute  structure  of  vegetables  as  re- 
vealed by  the  microscope,  i.  e.  to  the  composition  of  the  organs 
themselves.     But,  since  anatomy  in  the  animal  kingdom  includes 
the  consideration  of  general  as  well  as  of  minute  structure,  and 
indeed  answers  to  organography,  the  minute  anatomy  of  both 
kingdoms  takes  the  special  name  of  HISTOLOGY.     The  study  of 
functions,  or  of  the  living  being  (animal  or  plant)  in  action, 
is  the  province  of  PHYSIOLOGY. 

6.  SYSTEMATIC  BOTANY,  or  the  study  of  plants  in  their  kinds 
and  in  regard  to  their  relationships,  comprises  TAXONOMY,  or  the 
principles  of  classification,  as  derived  from  the  facts  and  ideas 


INTRODUCTION.  8 

upon  which  species,  genera,  &c.,  rest;  CLASSIFICATION  or  the 
SYSTEM  OF  PLANTS,  the  actual  arrangement  of  known  plants  in 
systematic  order  according  to  their  relationships  ;  PHYTOGRAPHT, 
the  rules  and  methods  of  describing  plants  ;  and  NOMENCLATURE, 
the  methods  and  rules  adopted  for  the  formation  of  botanical 
names.  GLOSSOLOGY  or  TERMINOLOGY1  is  a  necessary  part  of 
Phytography  or  Descriptive  Botany,  and  hardly  less  so  of 
Structural  Botany :  it  relates  to  the  application  of  distinctive 
terms  or  names  to  the  several  organs  or  parts  of  plants,  and  to 
their  numberless  modifications  of  form,  &c.  This  requires  a 
copious  vocabulary  of  well-defined  technical  terms,  by  the  use  of 
which  the  botanist  is  able  to  describe  the  objects  of  his  study 
with  a  precision  and  brevity  not  otherwise  attainable.  It  will 
be  convenient  to  exemplify  the  principal  terms  along  with  the 
modifications  of  conformation  which  they  designate ;  and  also, 
for  greater  fulness  and  facility  of  reference,  to  append  to  this 
volume  an  alphabetical  summary  of  them,  or  Vocabulary  of 
Botanical  Terms.2 

7.  The  present  volume  is  mainly  devoted  to  Morphological 
Botany ;   that  is,  to  Structural  Botany  on  the  basis  of  mor- 
phology.    This  department  cannot  be  properly  dealt  with  apart 
from  considerable  reference  to  intimate  structure,  development, 
and   function,  the   subject-matter   of  vegetable   histology  and 
physiology.     But  these  will  here  be  treated  only  in  the  most 
general  or  incidental  and  elementary  way,  and  only  so  far  as 
is  necessary  to   the  understanding  of  the  morphology  of  the 
stem,  leaves,  &c.     The  whole  discussion  of  the  histology  and 
physiology  of  plants  is  relegated  to  a  following  volume  and  to 
another  hand. 

8.  The  most  comprehensive  and  important  division  of  the 
vegetable  kingdom  is  into  plants  of  the  higher  and  of  the  lower 
series  or  grade,  i.e.  into  PH^NOGAMOUS  (or  PHANEROGAMOUS)  or 
FLOWERING,  and  CRYPTOGAMOUS  or  FLOWERLESS  PLANTS.     The 
first  are  all  manifest!}7  of  one  type,  and  therefore  have  a  consist- 
ent and  simple  morphology.     The  second  differ  among  them- 
selves almost  as  widely  as  they  do  from  the  higher  series  ;  and 


1  GLOSSOLOGY  is  the  better  word,  but  TERMINOLOGY,  although  a  hybrid 
of  Latin  and  Greek,  is  in  common  use. 

2  What  is  called  GEOGRAPHICAL  BOTANY  is  the  study  of  plants  in  respect 
to  their  natural  distribution  at  the  present  time  over  the  earth's  surface,  and 
the  causes  of  it.    FOSSIL  BOTANY  (Vegetable  Palaeontology)  relates  to  the 
plants  of  former  ages,  as  more  or  less  made  known  in  their  fossil  remains. 
MEDICAL  BOTANY,  AGRICULTURAL  BOTANY,  and  the  like,  are  applications 
of  Botany  to  medicine,  agriculture,  &c. 


4  INTRODUCTION. 

their  morphology  is  more  special  and  difficult.  Wherefore  it  is 
better  to  treat  them  separately  and  subsequently.  This  will  be 
done  in  a  third  part,  by  an  associate  devoted  to  Cryptogamic 
Botany. 

9.  Thus  the  field  is  here  left  clear  for  the  Structural  Botany 
of  Phaenogamous  or  Flowering  Plants,  with  which  the  study  of 
the  science  should  naturally  begin.  In  theory  it  may  seem 
proper  to  commence  with  the  simplest  plants  and  the  most  ele- 
mentary structures  ;  but  that  is  to  put  the  difficult  and  recondite 
before  the  plain  and  obvious.  The  type  or  plan  of  the  vegetable 
kingdom,  upon  which  morphological  botany  is  grounded,  is  fully 
exemplified  only  in  the  higher  grade  of  plants,  is  manifest  to 
simple  observation,  and  should  be  clearly  apprehended  at  the 
outset. 


CHAPTER  I. 

OUTLINES   OP  THE  GENEEAL   MORPHOLOGY  OF 
PH^ENOGAMOUS   PLANTS. 

10.  MORPHOLOGY,  the  doctrine  of  forms,  as  the  name  denotes, 
is  used  in  natural  history  in  nearly  the  same  sense  as  the  older 
term  Comparative  Anatomy.     If  it  were  concerned  merely  with 
the  description  and  classification  of  shapes  and  modifications, 
it  would  amount  to  little  more  than  glossology  and  organography. 
But  it  deals  with  these  from  a  peculiar  point  of  view,  and  under 
the  idea  of  unity  of  plan  or  type.1 

11.  As  all  vertebrate  animals  are  constructed  upon  one  type 
(or  ground  plan) ,  which  culminates  or  has  its  archetype  in  man, 
so  all  plants  of  the  higher  grade  (8)  are  strictly  of  one  type ; 
the  different  kinds  being  patterns  or  repetitions  of  it,  with  varia- 
tions.    The  vegetable  kingdom,  however,  does  not  culminate  in 
an  archetype  or  highest  representative.     As  respects  the  organs 
of  vegetation,  the  higher  classes  of  cryptogamous  plants  exhibit 
this  same  type ;   but  it  is  only  in  the  most  general  or  in  a 
recondite  sense  that  this  can  be  said  of  their  organs  of  repro- 
duction, and  of  the  less  differentiated  structure  of  the  lowest 
classes.     Wherefore   cryptogamous   plants   are  left  out  of  the 
present  view,  to  be  treated  apart. 

12.  Viewed  morphologically  and  as  to  its  component  organs, 
a  plant  is  seen  to  consist  of  an  axis  or  stem,  which  sends  off 
roots  into  the  soil,  and  bears  lateral  appendages,  commonly  as 
leaves,  but  which  may  be  very  unlike  leaves  in  whole  appearance 

1  The  term  Morphology  was  introduced  into  science  by  Goethe,  at  least  as 
early  as  the  year  1817  (Zur  Naturwissenschaft  iiberhaupt,  besonders  zur 
Morphologic,  Stuttgart  und  Tubingen,  1817-24).  On  page  9  of  the  first 
volume,  he  is  understood  to  have  suggested  this  word  for  the  purpose  and  in 
the  sense  now  adopted  in  botany  and  zoology.  It  essentially  replaces  an 
earlier  and  somewhat  misleading  word,  Metamorphosis,  (304.) 

Apparently  the  first  botanist  to  adopt  the  term  was  Auguste  de  St. 
Hilaire,  in  his  "  Lecons  de  Botanique,  comprenant  principalement  la  Mor- 
phologic Ve'ge'tale,  etc.,  Paris,  1841.  The  term  seems  not  to  have  been  taken 
up,  in  zoology,  by  Etienne  Geoffrey  Saint-Hilaire,  the  antagonist  of  Cuvier 
(who  was  of  a  wholly  different  family  from  that  of  the  botanist),  although 
the  game  idea  was  denoted  by  his  phrase  "  unity  of  organic  composition-" 


6  GENERAL  MORPHOLOGY 

and  function.  These  appendages,  whatever  their  form  or  use, 
accord  with  leaves  in  mode  of  origin,  position,  and  arrangement 
on  the  axis  or  stem.  Their  most  general  and  ordinary  form  is 
the  familiar  one  of  foliage  ;  hence  the  name  of  leaves  has  been 
by  botanists  extended  in  a  generic  way  from  the  green  expan- 
sions which  constitute  foliage  to  other  forms  under  which  such 
appendages  occur.  The  proper  morphological  expression  is, 
that  the  latter  are  homologous  with  leaves,  or  are  the  homologues 
of  leaves.1 

13.  Leaves  are  borne  upon  the  stem  at  definite  places,  which 
are  termed  NODES.     A  node  may  bear  a  single  leaf  or  a  greater 
number.     When  it  bears  two,  they  occupy  opposite  sides  of 
the  stem.     When  three,  four,  or  more,  they  divide  the  circum- 
ference of  the  stem  equally,    forming   a   circle,    technically  a 
WHORL,  or  in  Latin  form  a  VERTICIL.     When  only  two,  the  pair 
evidently  answers  to  the  simplest  kind  of  whorl.     So  that  leaves 
are  either  single  on  the   nodes,  in  which  case  they  are  alter- 
nate, that  is,  come  one  after  another  on  the  stem ;  or  in  whorls 
(whorled,  verticiUate) ,  in  the   commoner  case  of  a  single  pair 
being  called  opposite.     The  bare  space  between  two  successive 
nodes  is  an  INTERNODE.     This  is  longer  or  shorter,  according  to 
the  amount  of  longitudinal  growth,  which  thus  spaces  the  leaves, 
or  whorls  of  leaves,  in  most  various  degrees,  either  widely  when 
the  internodes  are  elongated,  or  slightly  when  they  remain  very 
short.     The  plant,  therefore  (roots  excepted) ,  is  made  up  of  a 
series  of  similar  parts,  i.  e.  of  portions  of  stem,  definitely  bearing 
leaves,  each  portion  developed  from  the  apex  of  the  preceding 
one.     This  constitutes  a  simple-stemmed  plant. 

14.  Branching  is  the  production  of  new  stems  from  the  older 
or  parent  stem.     These  normally  appear  in  the  AXILS  of  leaves, 
that  is,  in  the  upper  angle  which  the  leaf  forms  with  the  stem,  — 
from  which  they  grow  much  as  the  primary  stem  grew  from  the 
seed.     The  primary  stem,  connected  with  the  ground,  produces 
roots  which  develop  downwardly  into  the  soil,  from  which  they 
draw  sustenance.      Branches,   when  developed  above  ground, 

1  A  common  designation  for  all  these  appendages  being  desirable,  a  good 
one  is  furnished  by  the  Greek  name  for  leaf,  <pt\\ov,  PHYLLUM,  plural 
PHYLLA.  This,  used  with  prefixes,  may  be  made  to  designate  the  kind  of 
leaves  in  many  cases,  —  as,  prophylla,  catapky/la,  hypsophylla. 

Recent  German  botanists  use  the  word  Phyllome  in  this  sense.  It  is  a 
rather  convenient  and  well-sounding  word ;  but  phylloma  is  the  exact  Greek 
equivalent  of  our  word  foliage,  and  therefore  not  very  well  chosen  as  a 
common  term  for  leaves  which  are  not  foliage  as  well  as  those  which  are. 
Nor  will  this  word,  like  phyllum,  readily  take  prefixes,  as  above,  or  the  adjec- 
tive form,  as  it  readily  does  in  prophyllous,  hypsophyllous,  gamophyllous,  &c. 


OF   PELENOGAMOUS   PLANTS. 


being  in  organic  connection  with  their  parent  stem,  do  not 
usually  produce  roots ;  but  when  placed  in  equally  favorable 
conditions  for  it,  i.  e.  on  or  in  the  soil,  they  may  strike  root  as 
freely  as  does  the  original  stem. 

15.  An  incipient  stem  or  branch,  with  its  rudimentary  leaves, 
is  a  BUD.     The  normal  situation  of  a  bud  is  in  the  axil  of  a  leaf 
(axillary') ,  the  development  giving  rise  to  branches ;  or  else  at 
the  apex  of  an  axis  (terminal),  where  there  can  be  only  one,  the 
development  of  which  continues  that  axis.1 

16.  As  branches  are  repetitions  and  in  one  sense  progeny  of 
the  stem  which  bears  them,  so  the  serial  similar  parts  or  leaf- 
bearing  portions  of  a  simple  stem  are  repeti- 
tions, or  in  a  like  sense  progeny,  each  of  the 

preceding    one    from    which    it   grew.     The 
simple-stemmed  plant  is  made  up  of  a  series 
of  such  growths,  each  from  the   summit  of 
its  predecessor ;  the  branched  plant,  of  ad- 
ditional series,  laterally  developed,  from  ax- 
illary buds.     These  ultimate  similar  parts  into 
which   a   plant   may  thus   be  analyzed,  and 
which  are  endowed  with  or  may  produce  all 
the  fundamental  organs  of  vegetation,  were  by 
Gaudichaud    called    PHYTONS.      But   phyton, 
being  the  common  Greek  name  for  plant,  was 
not  a  happily  chosen  appellation   for   plant- 
elements,  or  homologous  plant-units 
term  for  them  is  PHYTOMERA 
peQog,  part),  equivalent  to  plant-parts 
structures  which,  produced  in  a  series,  make    ]/ 
up  a  plant  of  the  higher  grade.     In  English, 
the  singular  may  be  shortened  to  PHYTOMER. 

17.  This  theoretical  conception  of  the  organic 
composition  of  the  plant  is  practically  impor- 
tant to  the  correct  understanding  of  morpho- 
logical botany.  The  diagram,  Fig.  1,  serves 
to  represent  the  organic  elements,  orphytomera, 
in  a  simple  case,  such  as  that  of  a  growing 
plant  of  Indian  Corn,  or  other  Grass.  Here 

1  Bifurcation  by  the  division  of  a  terminal  bud  into  two,  as  in  Acrogenous 
Cryptogams,  is  supposed  by  some  to  occur,  even  normally,  in  some  Phaeno- 
gams,  especially  in  certain  forms  of  inflorescence ;  but  this  has  never  been 
convincingly  made  out. 

PIG.  1.  Diagram  of  a  simple-stemmed  plant,  exhibiting  the  similar  parta,  or 
phytomera,  a  to  h,  of  which  it  is  composed. 


on   for   plant- 
inits.   A  better  fft\\ 
(cpvrov,  plant,  V     \^y 
nt-parts,  —  the    \\      ^\ 


8  GENERAL  MORPHOLOGY 

the  leaves  are  alternate ;  in  other  words,  each  phytomer  is 
single-leaved ;  while  in  the  subsequent  illustrations  of  plants 
developed  from  the  seed,  at  least  the  earliest  phytomera  are 
two-leaved. 

18.  The  plan  thus  exhibited  in  the  leafy  stem  begins  in  the 
embryo,  or  initial  plant  in  the  seed,  and  is  carried  on  into  the 
flower,  in  which  the  normal  development  of  the  axis  finally  ends. 
One  plan  prevails  throughout.  To  illustrate  it,  the  morphology 
and  growth  of  the  embryo,  of  the  plant  developed  for  vegetation 
and  the  general  purposes  of  its  individual  existence,  and  lastly 
of  the  flower,  through  which  sexual  reproduction  takes  place, 
may  be  successively  treated  in  this  order. 


OF   THE  EMBRYO  AND   SEEDLING. 


CHAPTER  II. 

MORPHOLOGY   AND    DEVELOPMENT    OF    THE   EMBRYO   AND 
SEEDLING. 

19.  The  Embryo  is  the  initial  plant,  originated  in  the  seed.1 
In  some  seeds  it  is  so  simple  and  rudimentary  as  to  have  no 
visible  distinction  of  parts  :  in  others,  these  parts  may  have 
assumed  forms  which  disguise  their  proper  character.  But  every 
well-developed  embryo  essentially  consists  of  a  nascent  axis,  or 
stem,  bearing  at  one  end  a  nascent  leaf  or  leaves,  or  what  an- 
swers to  these,  while  from  the  other  and  naked  end  a  root  is 
normally  to  be  produced.  This  stem  is  the  primitive  internode 
of  the  plant :  its  leaf  or  pair  of  leaves  is  that  of  the  first  node. 
The  plant  therefore  begins  as  a  single  phytomer.  Some  embryos 
are  no  more  than  this,  even  when  they  have  completed  their 
proper  germination :  others  have  taken  a  further  development 
in  the  seed  itself,  and  exhibit  the  rudiments  of  one  or  more  fol- 
lowing phytomera.  The  embryo  of  the  Maple  is  an  example  of 
the  first  kind  ;  and,  being  large  enough  for  handling  and  for  the 
display  of  all  its  parts  to  the  naked  eye,  and  the  character  of 
these  parts  being  manifest  even  in  the  seed,  it  is  a  good  subject 
with  which  to  commence  this  study.  And  for  this  the  Sugar- 
Maple  is  one  of  the  best  of 
the  Maples.  Its  embryo 
(seen  in  Fig.  2  in  the  coiled 
condition  which  it  occupies 
in  the  seed,  and  in  Fig.  3 
and  Fig.  4  uncoiling  and  be- 
ginning to  grow)  is  an  initial  stem,  bearing  a  pair  of  leaves,  and 
nothing  more.  These  parts  take  the  technical  names  of 

1  Normally  a  seed  contains  a  single  embryo.  Polyembry,  the  formation  of 
two  or  more  embryos,  occurs  occasionally  as  a  kind  of  superfcetation  in 
some  seeds.  In  those  of  the  cultivated  Orange  it  is  most  common,  and  an 
evident  monstrosity.  In  Coniferse  and  Loranthacese,  two  or  three  embryos, 
of  equal  size  and  perfection,  are  not  rarely  produced. 

FIG.  2.  Embryo  of  Sugar  Maple,  in  vertical  section,  as  coiled  in  the  seed,  merely 
somewhat  loosened.  3.  Embryo  of  same,  just  beginning  to  unfold  in  germination. 
4.  Same  more  advanced  :  a.  its  stem  or  caulicle ;  bb.  its  two  leaves  or  cotyledons. 


10 


MORPHOLOGY 


20.  Caulicle  or  Radicle,  and  Cotyledons.  The  name  of  radicle 
was  early  applied  to  the  axis  of  the  embryo  below  the  cotyledons, 
on  the  supposition  that  it  was  the  actual  beginning  of  the  root. 
But  its  structure  and  mode  of  growth  show  it  is  not  root  (24, 
44,  78),  but  a  body  of  the  exact  nature  of  stem,  from  the 
naked  end  of  which  the  root  is  developed.  Wherefore  Caulicle 
(Lat.  cauliculus,  diminutive  of  caul-is,  stem)  is 
the  appropriate  name ;  and  it  would  be  gen- 
erally adopted,  were  it  not  that  the  older  term 
is  so  incorporated  into  the  language  of  sys- 
tematic botany  (in  which  fixity  and  uniformity 
are  of  the  utmost  importance)  that  it  is  not 
easily  displaced.  It  may  be  continued  in 
descriptive  botany  on  this  account,  but  in 
morphology  it  is  apt  to  mislead  ;  and  the  name 
of  caulicle,  suggestive  of  the  true  nature  of 
the  organ,  is  preferable.1  The  more  fanciful 
name  of  Cotyledons  was  very  early  applied  to 
what  are  now  recognized  as  answering  to  the 
leaves  of  the  embryo  :  it  has  the  negative  merit 
of  suggesting  no  misleading  analogy.2 

21.  Development  of  the  Dicotyledonous  Em- 
bryo, i.  e.  the  two-leaved  embryo.  This,  in 
the  Red  Maple  (Figs.  5-8),  usually  germinates 
in  summer,  shortly  after  the  fruits  of  the  season 
have  matured  and  fallen  to  the  ground.  It 
differs  from  that  of  Sugar  Maple  in  the  crump- 
ling instead  of  coiling  of  the  cotyledons  in  the 
seed.  Referring  the  whole  physiology  of  ger- 
mination to  that  part  of  the  work  which  treats 
of  Vegetable  Physiology,  the  development  of 
the  embryo  into  the  seedling  may  here  be  described,  taking  that  of 
a  Maple  for  a  convenient  type  or  pattern,  with  which  other  forms 


1  Linnaeus  called  it  RosteUum,  a  name  which,  being  etymologically  mean- 
ingless in  this  connection,  is  not  misleading.    The  French  botanists  named  it 
Tigelle,  diminutive  of  tige,  stem:  but  some  (like  Mirbel)  applied  the  term  to 
the  developing  axis  above  the  cotyledons;  others,  to  the  early  axis  both 
above  and  below  them.    The  name  Radicula  originated  with  Gaertner. 

2  The  name  Cotyledon,  which  was  adopted  by  Linnaeus,  is  a  Greek  word 
for  a  cup-shaped  hollow  or  cavity,  also  for  a  plant  with  thickish  and  saucer- 
shaped  leaves.     It  was  primarily  applied  to  the  thickened  "  lobes  "  of  the 
embryo,  the  foliaceous  nature  of  which  was  not  recognized. 

FIG.  6.  One  of  the  twin  winged  fruits  of  Red  Maple  (Acer  rubrum),  with  body 
divided,  to  show  the  seed.  6.  Seed  extracted  and  divided,  to  show  the  embryo  within. 
7.  Embryo  partlv  unfolded.  8.  Embryo  in  early  stage  of  germination. 


OF  THE  EMBRYO  AND  SEEDLING.  11 

may  afterward  be  compared.  The  first  growth  is  seen  in  the 
elongation  of  the  radicle  or  caulicle,  and  its  assumption,  as  far  as 
possible,  of  a  vertical  position,  and  the  production  of  a  root  from 
the  naked  end.  As  it  emerges  from  the  seed  in  consequence  of 
this  elongation,  the  root-end  of  the  caulicle  points  downward  into 
the  soil,  the  caulicle  bending,  if  need  be,  to  assume  this  position  ; 
and  the  nascent  root,  partaking  of  this  disposition,  grows  in  a 
downward  direction.  Hence  the  root  has  been  called  the  Descend- 
ing Axis  of  the  plant.  While  this  avoids,  the  opposite  or  budding 
end  (as  it  may  be  termed)  seeks  the  light,  and  when  free  takes 
an  upward  direction.  The  result  of  this,  and  of  the  elongation 
of  the  caulicle,  is  to  carry  the  budding  end  out  of  the  soil  and 
into  the  air,  where  the  growing  cotyledons  unfold  or  expand  and 
become  the  first  leaves,  or  Seed-leaves.  This  initial  stem  and  its 
continuation  therefore  constitutes  the  Ascending  Axis.  If  the 
budding  end  happen  to  lie  pointing  downward  and  the  root-end 
upward  in  the  ground  when  germination  begins,  both  will  curve 
quite  round,  as  they  grow,  to  assume  their  appropriate  directions. 
If  obstacles  intervene,  each  will  take  as  nearly  as  possible  its 
wonted  direction,  through  an  instinctive  tendency  and  action, 
which  insures  that  each  part  of  the  plant  shall  be  developed  in 
its  fit  medium,  —  the  root  in  the  dark  and  moist  earth,  the  stem 
and  leaves  in  the  light  and  air. 

22.  The  plantlet,  thus  established,  has  now  all  the  essential 
Organs  of  Vegetation,  as  they  are  called, 

i.  e.  root,  stem,  and  leaves.  Its  subse- 
quent development,  so  far  as  vegetation 
(apart  from  proper  reproduction)  is  con- 
cerned, consists  in  the  addition  of  more 
of  these,  until  the  whole  herb,  shrub,  or 
tree  is  built  up. 

23.  In  Maples  (as  in  the  Morning  Glory, 
Fig.  16,  and  many  others)  the  embryo  in 
the  seed,  and  until  after  the  full  develop- 
ment  of   its   cotyledons   or   seed-leaves, 
shows   no  rudiments  of  the   subsequent 
growth.    The  embryo  grows  into  the  plant- 
let  wholly  by  the  appropriation  of  prepared 
nourishing  matter  which  was  provided  by 

the  mother-plant  and  stored  in  the  seed, — in  the  case  of  the 
Maple,  wholly  in  the  embryo  itself,  mainly  in  its  cotyledons. 

FIG.  9.  Maple  embryo  developed  into  plantlet  of  one  phytomer,  and  producing 
rudiments  of  the  second:  the  lower  portion  covered  with  root-hairs  is  the  root;  the 
naked  portion  above  is  the  caulicle. 


12 


MORPHOLOGY 


After  this  is  consumed  and  in  good  part  converted  into  struc- 
ture, the  plantlet  must  by  the  action  of  its  root  and  leaves  imbibe 
from  the  soil  and  air  appropriate  materials,  and 
assimilate  them  into  nourishing  matter  needful 
for  further  growth.  Only  then  does  the  rudi- 
ment of  new  structure  appear,  in  the  form  of  a 
growing  point,  or  bud,  at  the  node  or  apex  of 
the  primitive  stemlet,  between  the  two  seed- 
leaves.  In  this  case  it  soon  shows  itself  as 
a  second  pair  of  leaves,  at  first  resting  on 
the  node  (Fig.  9),  next  as  somewhat  upraised 
by  the  development  of  the  second  internode 
(Fig.  10,  summit),  and  finally  both  this  inter- 
node  and  the  pair  of  leaves  complete  their 
growth  (Fig.  11).  Then  the  terminal  bud 
which  crowns  the  second  node  develops  in 
the  same  way  the  third  pair  of  leaves  and 
their  supporting  internode  or  joint  of  stem 
(Fig.  12)  ;  and  so  on. 

24.  The  root  and  the  stem  grow  not  only 
in  opposite  directions,  but  in  a  different  mode. 
The  primordial  stem,  pre-existing  in  the  seed 
(though  at  first  it  may  be  extremely  short) 
grows  throughout  its  whole  length,  but  most 
in  its  upper  part,  so  that  it  may  become  a 
stemlet  two  or  three  inches  long.  But,  soon 
attaining  its  full  growth  as  to  length,  the 
stem  is  carried  upwards  by  the  subsequent  joints  or  portions, 
similarly  developed  and  elongated,  one  after  the  other.  Not 
that  each  portion  necessarily  waits  until  the  growth  of  its  prede- 
cessor is  complete,  —  though  this  occurs  at  first  in  seedling  Maples 
and  other  embryos  unprovided  with  much  store  of  food,  — yet  the 
development  follows  this  course  and  order  of  succession.  The 
root,  on  the  contrary,  cannot  be  said  to  pre-exist  in  the  seed,  or 
at  most  it  may  be  said  to  exist  potentially  in  tissue  of  the  caulicle 
from  which  a  root  or  roots  normally  originate.1  It  is  formed 

1  Yet  from  nothing  which  is  special  to  this  part  of  the  embryo,  nor  to  the 
embryo  at  all.  The  primary  root  is  developed  from  subjacent  tissue  of  the 
tip  of  the  caulicle,  just  as  it  is  sometimes  developed  from  along  the  sides, 
and  as  secondary  roots  are  from  all  or  most  stems  under  favoring  conditions. 
This  complete  similarity,  and  the  fact  of  what  is  called  the  "  endogenous  " 
origin  of  roots  (i.  e.  their  springing  from  subjacent  rather  than  superficial 
tissue)  appear  fully  to  warrant  the  statement  in  the  text  above. 

FIG.  10.  Maple  plantlet  with  second  internode  developing.  11.  Same  with  second 
internode  and  pair  of  leaves  complete,  and  bud  of  the  third  apparent. 


OP  THE  EMBRYO  AND   SEEDLING. 


13 


in  the  process  of  germination,  and  originates  in  tissue  just  back 
of  that  which  covers  the  root-end  of  the  caulicle,  and  which, 
being  carried  forward  by  the  subjacent 
formation  (to  which  it  becomes  a  sort  of 
cap  or  sheath) ,  is  called  the  Root-cap. 
As  the  primary  root  thus  began  by  a 
new  and  local  growth  at  the  extremity 
of  pre-existing  stem,  so  it  goes  on  to 
grow  in  length  wholly  or  mainly  by  a 
continuation  of  this  formation,  the  new 
at  the  end  of  the  old.  That  is,  the 
root  elongates  by  continual  minute 
increment  of  its  apex  or  near  it,  the 
formed  parts  very  soon  ceasing  to 
lengthen.  This  is  in  marked  distinc- 
tion from  stem,  which  grows  by  suc- 
cessive individualized  portions ;  and 
these  portions  (internodes) ,  at  first 
very  short,  attain  or  are  capable  of 
attaining  a  considerable  and  sometimes 
very  great,  but  definitely  terminable 
length,  by  interstitial  growth  through- 
out. Moreover,  roots  are  naked,  not 
producing  as  they  grow  either  leaves 
or  any  organs  homologous  with  leaves. 
They  commonly  branch  or  divide,  but 
in  a  vague  manner ;  and  their  new  parts  bear  what  are  called 
Root-hairs,  which  greatly  increase  the  absorbing  surface  ;  other- 
wise they  are  destitute  of  appendages  or  organs. 

25.  With  the  Maple  embryo,  here  taken  as  a  type,  that  oi 
Morning  Glory,  Ipomcea  purpurea,  or  any  of  its  kin,  may  next 
be  compared.  The  cotyledons  are  different  in  shape,  being  as 
broad  as  long,  and  notched  both  at  base  and  apex.  They  lie 
in  contact  in  Fig.  14,  and  are  very  thin,  leaf-like,  and  green 
while  contained  in  the  seed.  Their  thinness  is  shown  in  Fig.  13, 
where  a  section  of  the  crumpled  and  folded  embryo,  as  it  lies  in 
the  seed,  exactly  divides  them  (passing  through  the  terminal  and 
basal  notches)  and  also  the  caulicle,  which  here  is  thicker  than 
both.  The  germination  is  similar  to  that  of  the  Maple  ;  and  like 
that  (as  Fig.  16  shows),  and  for  the  same  reason,  no  bud  or 
rudiment  of  the  further  growth  pre-exists  in  the  embryo  or 


FIG.  12.    Red  Maple  seedling,  with  three  joints  of  stem  and  pairs  of  leave*  developed, 
the  first  being  the  cotyledons. 


14 


MORPHOLOGY 


appears  in  the  young  plantlet,  until  that  has  established  itself 
and  had  time  to  elaborate  proper  material  therefor.  This  con- 
dition is  correlated  with  thin  foliaceous 
cotyledons,  holding  no  store  of  nourish- 
ment. Here  they  do  not  contain  sufficient 
material  for  the  development  of  the  initial 
stem  and  root.  The  maternal  provision 
for  this  is  here  stored  up  in  the  seed 
around  but  not  within  the  embryo.  This 
nourishing  deposit,  seen  in  the  section 
(Fig.  13)  filling  the  whole  space  between 
the  seed-coats  and  the  thin  embryo,  was 
named  by  the  early  botanists  and  vege- 
table anatomists  the  ALBUMEN  of  the  seed.1 
This  substance,  softened  in  germination 
and  by  chemical  changes  rendered  soluble, 
is  gradually  absorbed  by  the  cotyledons 
as  material  for  their  growth  and  that  of 
the  developing  primary  stem  and  root. 

26.  Seeds  in  this  regard  are  accordingly 
distinguished  into  albuminous  and  exal- 
buminous,  those  supplied  with  and  those 
destitute  of  albumen.  The  difference 
inheres  neither  in  the  character  nor  in 
the  amount  of  the  maternal  provision  for  the  development  of 
the  embryo-plant,  but  merely  in  the  storage.  In  exalbuminous 
seeds  the  nourishment  supplied  for  this  purpose  is  taken  into 
the  embryo  itself,  mostly  into  the  cotyledons,  during  the  growth 
and  before  the  maturity  of  the  seed.  In  albuminous  seeds 
this  same  material  is  deposited  around  or  at  least  external  to 
the  embryo. 

27.  The  amount  of  this  deposit  is,  in  the  main,  inversely  pro- 

1  Grew  appears  to  have  first  applied  this  name,  and  Gaertner  to  have 
introduced  it  into  systematic  botany,  where  it  remains  in  use,  although 
Jussieu  replaced  it  by  the  term  Perisperm,  and  Richard  by  Endosperm, 
neither  of  them  much  better  etymologically  than  the  old  word  Albumen. 
But  it  must  be  kept  in  mind  that  it  was  intended  to  liken  the  "  albumen  "  of 
the  seed  with  the  albumen  or  white  of  an  egg  as  a  body  or  mass,  and  not  as 
a  chemical  substance ;  the  embryo  being  fancifully  conceived  to  be  analo- 
gous to  the  yolk  of  the  egg,  the  surrounding  substance  of  this  kind  not 
unnaturally  took  the  name  of  the  white,  viz.  albumen. 

FIG.  13.  Section  of  seed  of  common  Morning  Glory,  Ipomoea  purpurea,  dividing 
the  contained  embryo  through  the  centre.  14.  Embryo  of  same,  detached  and  straight- 
ened. 15.  Embryo  in  germination ;  the  cotyledons  only  partly  detached  from  the  coat 
of  the  seed.  16.  Same,  later  and  more  developed,  the  cotyledons  unfolded  and  out- 
tpread  as  the  first  pair  of  leaves. 


OF   THE   EMBRYO   AND   SEEDLING. 


15 


portional  to  the  size  and  strength  of  the  embryo,  or  the  degree 
of  its  development  in  the  seed.  A  comparison  of  the  various 
illustrations  sufficiently  shows  this.  Figures  17  to  24  exhibit, 
in  a  few  common 
seeds,  somewhat  of 
this  relation,  and 
also  of  the  position 
and  shape  assumed 
in  some  instances. 


The  upper  rank  of 
figures  represents 
sections  of  seeds ; 
the  embryo  left  in 
white ;  the  albumen 
as  a  dotted  surface. 
The  lower  rank  shows  the  embryos  detached.  That  of  Mirabilis 
has  very  broad  and  thin  cotyledons,  a  caulicle  of  equal  length, 
and  the  whole  curved  round  the  albumen  which  thus  occupies 
the  centre  of  the  seed.  That  of  Potato  is  coiled  in  the  midst 
of  the  albumen,  is  slender ;  the  cotyledons  narrowed  down  to 
semi-cylindrical  bodies,  not  leaf-like  in  appearance,  and  the  two 
together  not  thicker  than  the  caulicle.  In  Barberry  the  embryo 
is  straight,  in  the  axis  of  the  albumen,  which  it  almost  equals  in 
length ;  the  cotyledons  considerably  broader  than  the  caulicle, 
but  short  and  thickish.  That  of  the  Peony  is  similar,  but  very 
much  smaller,  occupying  a  small  space  at  one  end  of  the  albu- 
men, and  seemingly  without  distinction  of  parts,  but  under  the 
microscope  and  with  some  manipulation  the  broader  end  is 
found  to  be  divided,  that  is,  to  consist  of  two  minute  cotyledons. 
The  embryo  of  a  Crowfoot  is  similar,  but  still  more  minute  and 
the  parts  hardly  to  be  distinguished ;  and  in  some  minute  em- 
bryos there  is  no  apparent  distinction  of  parts  until  they  develop 
in  germination. " 

28.  The  study  of  the  formation  of  the  embryo  in  the  seed 
teaches  that  all  embryos  begin  with  a  still  more  simple,  minute, 
and  homogeneous  structure ;  and  these  comparisons  suffice  to 
show  that  all  such  differences  are  referable  to  different  degrees 
and  somewhat  different  modes  of  the  development  of  the  embryo 
while  yet  in  the  seed.  It  also  appears  that  the  size  and  shape 


FIG.  17.     Section   of  seed    and   contained    embryo    of  Mirabilis   (Four-o-clock). 
18.  Embryo  detached  entire. 

FIG.  19.  Section  of  a  Potato-seed.  20.  Embryo  detached  entire. 
FIG.  21.  Section  of  Barberry-seed.  22.  Embryo  detached  entire. 
FIG.  23.  Section  of  Peony-seed.  24.  Embryo  detached  entire. 


16 


MOEPHOLOGY 


of  an  organ  do  not  indicate  its  nature,  either  in  the  embryo  or 
in  subsequent  growth.  But  in  all  the  cases  yet  mentioned  the 
cotyledons  actually  demonstrate  their 
nature  by  developing  in  germination 
in  a  foliaceous  manner  and  becoming 
the  first  leaves  of  the  seedling.  Nor 
is  this  nature  much  disguised  by  the 
fact  that  the}'  differ  greatly  in  form  in 
different  species,  and  that  the  seed- 
leaves,  or  developed  cotyledons,  differ 
much  in  shape  and  often  in  texture 
from  the  succeeding  leaves.  (See  Fig. 
11,  12,  25,  &c.) 

29.  To  complete  the  comparison 
between  the  seedling  Morning  Glory 
and  that  of  the  Maple,  it  is  to  be 
noted  that  here,  while  the  cotyledons 
or  seed-leaves  are  two,  the  following 
internode  bears  only  one  leaf  (Fig.  25),  as  also  will  the  just  de- 
veloping third  internode ;  and  this  continues  throughout  up  to 
the  blossom :  that  is,  the  leaves  subsequent 
to  the  cotyledons  are  not  opposite  as  in  the 
Maple,  but  alternate.  (13.) 

30.  All  the  preceding  illustrations  are  from 
embryos  which  previous  to  germination  have 
developed  nothing  beyond  the  cotyledons.  In 
the  following,  a  rudiment  of  further  growth, 


26  2T  28  29 

or  a  primary  terminal  bud,  is  visible  in  the  seed.     It  is  most 
manifest  in  large  and  strong  embryos  with  thick  or  flesh}*  cotyle- 

FIQ.  25.  Further  development  of  Morning  Glory,  Fig.  16,  the  root  cut  away,  the 
internode  above  the  cotyledons  and  its  leaf  completed,  the  next  internode  and  its  leaf 
appearing. 

FIG.  26.  Embryo  (kernel)  of  the  Almond.  27.  Same,  with  one  cotyledon  removed, 
to  show  the  plumule,  a. 

FIG.  28.  Section  of  an  Apple-seed,  magnified,  cutting  through  the  thickness  of  the 
cotyledons.  29.  Embryo  of  the  same,  extracted  entire,  the  cotyledons  a  little  separated, 

FIG.  30.  Germination  of  the  Cherry,  showing  the  thick  cotyledons  little  altered, 
and  the  plumule  developing  the  earliest  real  foliage. 


OF   THE  EMBRYO  AND   SEEDLING. 


17 


dons,  t.  e.  cotyledons  well  charged  with  nourishing  matter.  The 
early  vegetable  physiologists  gave  to  it  the  name  of  PLUMULE 
(Lat.  plumula,  a  little  plume).  The 
name  was  suggested  by  its  appearance 
in  such  an  embryo  as  that  of  the  bean 
(Phaseolus) ,  in  which  it  evidently  con- 
sists of  a  rudimentary  pair  of  leaves, 
while  in  the  pea  and  the  acorn  it  is  a 
rudimentar}*  stem,  the  leaves  of  which 
appear  only  later,  when  germination 
has  considerably  advanced.  In  any 
case,  the  plumule  is  the  bud  of  the 
ascending  axis  already  discernible  in 
the  seed.  Fig.  27,  a,  shows  it  in  the 
almond,  one  cotyledon  being  removed. 
Fig.  28  shows  it  in  the  section  of  a 
similar  although  much  smaller  embryo, 
that  of  an  apple-seed,  enlarged  to 
nearly  the  size  of  the  other.  It  is 
equally  visible  in  the  cherry,  the  bean, 
and  the  beechnut.  The  embryo  in  all 
these  cases  constitutes  the  whole  kernel 
of  the  seed.  For  the  nourishment, 
which  in  all  the  foregoing  illustrations 
except  the  first  (i.e.  in  Fig.  13, 17-23), 
is  deposited  around  or  exterior  to  the 
embryo,  is  in  these  stored  within  it. 

31.  The  development  of  these  em- 
bryos in  germination  proceeds  in  the 
normal  manner,  but  with  two  cor- 
related peculiarities.  First,  by  the 
lengthening  of  the  radicle  more  or  less,  their  thick  cotyledons 
are  usually  raised  to  or  above  the  surface  of  the  soil ;  they 
expand,  assume  the  green  color  needful  to  foliage ;  but  they 
imperfectl}'  or  in  a  small  degree  perform  the  function  of 
green  leaves.  Their  main  office  is  to  suppty  the  other  growing 
parts  with  the  prepared  nourishment  which  they  abundantly 
contain.  Then,  being  thus  copiously  nourished,  the  root  below 
and  the  ready-formed  plumule  above  grow  rapidly  and  strongly, 
having  accumulated  capital  to  draw  upon  ;  and  the  leaves  of  the 


FIG.  31.     Beechnut  cnt  across,  filled  by  the  fleshy  embryo;  the  thick  cotyledon* 
partly  enfolding  each  other.    32.  Embryo  of  the  same  in  early  germination     33.  Same 
more  advanced;  the  plumule,  which  is  just  emerging  in  the  preceding,  here  developed 
Into  a  long  internode  and  a  pair  of  leaves. 
5V 


18 


MORPHOLOGY 


latter  are  practically  the  earliest  efficient  foliage  of  the  plantlet. 
Thus,  as  in  the  germinating  Cherry-seed  (Fig.  30) ,  three  or  four 
internodes  of  stem,  with  their  leaves,  may  be  produced  before 
these  leaves  themselves  are  sufficiently  developed  to  make  any 
sensible  contribution  to  this  growth.  And  in  the  Beech  and  Bean, 


the  leaves  of  the  plumule  come  forward  almost  before  the  root 
has  attached  the  plantlet  to  the  soil.  (Fig.  32,  35.)  Between 
such  cases  and  that  of  Maple  and  the  like  there  are  all  degrees. 
There  are  also  familiar  cases  in  which  the  storage  of  nourishment 
in  the  cotyledons  is  carried  to  a  maximum,  with  results  which 
gravely  affect  the  development. 


PIG.  34.  The  embryo  (the  whole  kernel)  of  the  Bean.  35.  Same  early  in  germi- 
nation; the  thick  cotyledons  expanding  and  showing  the  plumule.  36.  Same,  more 
advanced  in  germination  ;  the  plumule  developed  into  an  internode  of  stem  bearing  a 
pair  of  leaves. 

FIG.  37.  Embryo  of  Pea,  i.  e,  a  pea  minus  the  seed-coat.  38.  Advanced  germi- 
nation of  the  same. 


OP   THE   EMBRYO   AND   SEEDLING. 


19 


32.  Thus,  in  the  Pea,  near  relative  of  the  Bean,  the  embryo 

(Fig.  37),  which  is  the  whole  kernel  of  the   seed,  has  the 

cotyledons  so  gorged  with  this  nutritive 

store  that   they  are   hemispherical ;   and 

the   acorn  of  the   Oak   (Fig.  39),  near 

relative  of  the  Beech,  is  in  similar  case. 

These  extremely  obese  cotyledons  have 

not  only  lost  all  likeness  to  leaves,  but  all 

power  of  fulfilling  the  office  of  foliage, 

which  is  apparently  no  disadvantage  ;  for 

when  two  different  duties  are  performed 

by  the  same  organ,  it  rarely  performs  both 

equally  well.      Here  they  become  mere 

receptacles  of  prepared  food-,  the  nature 

and  office  of  which  is  the  same  as  of  the 

albumen,  or  nutritive  deposit  exterior  to 

the  embrj-o  in  what  are  called  albuminous 

seeds.  (25-27.)     The  difference  is  in  the 

place  rather  than  in  the  character  of  the« 

deposit.      The  plumule  in  such  cases  is 

always  apparent  before  germination  ;  and 
it  develops  even  with  more  vigor  than  in 
the  preceding  cases.  It  usually  rises  as  a 
stout  stem  of  several  internodes  lengthen- 
ing almost  simultaneously,  or  at  least  the 
upper  strongly  developing  long  before  the 
lower  have  finished  their  growth ;  and 
the  latter  are  practically  leafless,  bearing 
only  small  and  scale-like  and  useless  ru- 
diments of  leaves.  This  is  correlated  with 
the  peculiarity  that  the  caulicle  does  not 
lengthen  in  germination,  or  it  lengthens 
very  slightly ;  the  cotyledons  remain  within 
the  coats  of  the  seed ;  and  if  this  were 
buried  beneath  the  surface  of  the  ground,  there  it  remains.  The 
abortion  of  the  earliest  leaves  of  the  plumule  is  in  correlation 
with  this  hypogceous  (i.  e.  underground)  situation  of  the  cotyle- 
dons throughout  the  germination.  The  slight  elongation  of  the 
caulicle  serves  merely  to  protrude  its  root-end  from  the  coats  of 
the  seed  in  a  downward  direction,  and  from  this  a  strong  root 
usually  is  formed. 


FIG.  39.    Section  of  an  acorn,  filled  by  the  embryo.    40.  Advanced  germination  ot 
the  same. 


20 


MORPHOLOGY 


33.  In  some  Oaks,  notably  in  our  Live  Oak  (Quercus  virens), 
and  less  so  in  the  Horsechestnut,  the  two  cotyledons  coalesce  or 
cohere  by  their  contiguous  faces. 
In  some  of  these  cases  of  hypo- 
gaeous  germination,  the  short 
caulicle  and  plumule  are  extri- 
cated from  the  enclosing  coats  or 
husk  by  the  development  of  short 
stalks  (petioles,  157)  to  the  fleshy 
cotyledons  ;  as  is  seen  in  Fig.  42, 
and  in  most  germinating  acorns. 
These  petioles  are  not  visible  in 
the  seed,  but  are  the  first  develop- 
ment in  germination. 

34.  There  are  some  curious 
cases  in  which,  while  the  caulicle 
remains  short  and  subterranean, 
the  cotyledons  are  raised  out  of 
ground  in  germination  by  the 
formation  of  far  longer  stalks 
(petioles)  than  those  of  the 
Horsechestnut.  A  singularly  dis- 
guised instance  of  this  kind  is  seen  in  Megarrhiza,  a  genus  of 
Cucurbitaceous  plants  of  California  and  Oregon,  remarkable  for 
their  huge  root.  The  large  seed  has  very  thick  and  fleshy 
cotyledons,  and  a  very  short  and  straight  caulicle.  In  germi- 
nation, the  whole  seed  is  elevated,  seemingly  in  the  manner  of 
the  bean,  upon  a  stout  stem.  One  waits  for  a  long  time  expect- 
ing to  see  the  cotyledons  throw  off  the  bursting  husk  and  expand, 
or  else  to  put  forth  the  plumule  from  between  their  bases.  But 
at  length  the  plumule  makes  its  appearance  from  an  unexpected 
place,  coming  separately  out  of  the  soil.  Eemoving  this,  the 
state  of  things  represented  in  Fig.  43  is  presented, — that  of 
the  plumule  seemingly  originating  from  the  base,  instead  of  the 
apex,  of  an  elongated  caulicle  !  But  on  examination  of  the  cleft 
from  which  this  proceeds,  by  making  a  section  of  the  stem  above 
(showing  that  it  is  hollow),  and  finally  by  separating  the  cotyle- 
dons and  gently  tearing  apart  the  two  short  stalks  by  which  they 
are  united  to  their  stem-like  support,  it  is  found  that  the  latter  may 
be  divided  into  two  (as  shown  in  Fig.  44),  even  down  to  the  cleft 
below.  This  explains  the  anomaly.  The  real  caulicle  has  re- 

FIG.  41.  Section  of  a  Horsechestnut  or  Buckeye  seed,  through  the  very  thick 
cotyledons  and  the  incurved  caulicle.  42.  Seed  In  germination,  showing  the  petioles 
to  the  cotyledons,  <fcc. 


OF   THE  EMBRYO  AND   SEEDLING. 


21 


mained  short  and  subterranean,  and  is  confluent  with  the  upper 
part  of  the  thickening  root :  the  seeming  caulicle,  which  raised 
the  cotyledons  above  the  soil,  consists 
of  the  petioles  of  these  combined  into 
a  tubular  stem-like  body,  no  evident 
trace  of  which  is  visible  in  the  seed, 
although  in  germination  it  attains  the 
length  of  two  or  three  inches  :  in  age 
it  is  readily  separable  into  the  two 
leaf-stalks  or  petioles  of  which  it  is 
composed  :  the  plumule  is  thus  seen 
to  be  wholly  normal,  originating  from 
between  the  cotyledons.  All  the  ex- 
tensive growth  so  far,  and  until  the 
proper  foliage-leaves  of  the  continu- 
ation of  the  plumule  are  developed 
and  begin  their  action,  is  from  nutri- 
tive material  stored  in  the  thickened 
cotyledons,  a  considerable  part  of 
which  was  transferred  to  the  already 
enlarging  root,  before  a  remaining 
portion  was  used  in  building  up  the 
strong  plumule.  The  economy  of  this 
elevation  of  cotyle- 
dons which  never 
open,  and  of  the 
lengthened  distance 
through  which  the 
nutritive  matter  has 
to  be  carried,  is  not 
apparent.  But  it  is 
the  family  habit  in 
Cucurbitacese  to 
bring  up  the  cotyle- 
dons that  they  may 
develop  as  leaves 
(as  in  the  Pumpkin, 
Fig.  47) :  here  this 

elevation  is  brought  44  43 

about  in  a  different  way,  but  without  securing  the  useful  end.1 

1  It  may  be  inferred  that  Megarrhiza  is  a  descendant  of  some  Cucurbitacea 
with  thinner  cotyledons,  which  in  germination  developed  into  long-stalked 
leaves,  in  the  manner  described  in  the  next  following  paragraphs. 

FIG.  43,  44.    Peculiar  germination  of  Megarrhiza  Californica;  explained  above. 


22  MORPHOLOGY 

35.  This  same  anomaly,  as  to  the  development  of  long  stalks 
to  the  cotyledons  and  their  union  into  a  stem-like  body,  occurs  in 
various  species  of  Larkspur  (notably  in  the  Californian  Delphin- 
ium nudicaule)  ;  but  in  these  the  cotyledons  develop  into  a  pair 
of  efficient  green  leaves. 


45 

36.  A  similar  elongation  of  petioles  of  the  cotyledons,  but 
without  any  union,  occurs  in  a  species  of  Morning  Glory  of  the 
plains  beyond  the  Mississippi  (Ipomcaa  leptophylla)  ;  the  leaf- 
like  cotyledons  coming  up  on  their  long  stalks  separately  from 
the  ground    (Fig.  45)  ;   the   developed   plumule    rising   some 
time  afterward  between  them.     Compare  this  with  the  ordinary 
species  (25,  Fig.  15, 16,  25),  and  note  that  the  difference  is  merely 
that  the  caulicle  in  the  common  Morning  Glory  elongates  and  the 
petioles  of  the  cotyledons  remain  short. 

37.  In  all  instances  thus  far  a  single  primary  root  so  regularly 
develops  from  the  lower  end  of  the  axis  of  the  embryo  (variously 
named  radicle  or  caulicle) ,  and  forms  such  a  direct  downward 

FIG.  45.  Germination  of  Ipomoea  leptophylla;  the  caulicle  not  developing,  the 
plumule  and  the  petioled  cotyledons  rise  from  underground.  Dotted  line  marks  the 
level  of  the  soil. 

FIG.  46.  Embryo  of  a  Pumpkin,  the  cotyledons  separated.  47.  Same  germinated; 
a  cluster  of  roots  from  the  base  of  caulicle.  . 


OF  THE  EMBKYO  AND   SEEDLING.  23 

prolongation  of  it,  that  it  was  called  the  descending  axis ;  and 
the  body  from  which  it  originates  was  named  the  radicle,  on  the 
supposition  that  it  was  itself  the  nascent  root.  But,  as  already 
explained,  the  so-called  radicle  grows  in  the  manner  of  stem  (24), 
and  is  morphologically  that  initial  internode  the  node  of  which 
bears  the  first  leaves  or  cotyledons.  (20.)  Let  it  now  be  noted 
that  this  descending  axis  or  single  primary  root  is  far  from 
universal.  In  Pumpkin,  Squash,  Echinocystis,  and  the  like, 
the  strong  caulicle  sends  out  directly  from  its  root-end  a  cluster 
of  roots  or  rootlets,  of  equal  strength;  *.  e.,  it  strikes  root  in 
nearly  the  manner  that  a  cutting  does.  (Fig.  47.) 

38.  The  Polycotyledonous  Embryo  is  one  having  a  whorl  of 
more  than  two  seed-leaves.     The  dicotyledonous  embryo  being 
a  whorl  of  the  very  simplest  kind,  that  is,  with  the 

members  reduced  to  two,  the  polycotyledonous 
may  be  regarded  as  a  variation  of  it.  In  all  but 
one  group  of  plants  it  is  simply  a  variation,  of 
casual  occurrence,  or  even  a  monstrosity,  in  which 
three  or  rarely  four  cotyledons  appear  instead  of 
two.  In  Pines  (Fig.  48,  49),  however,  and  in 
most  but  not  all  Coniferse,  a  whorl  of  from  3  to  10 
cotyledons  is  the  normal  structure,  varying  accord- 
ing to  the  species,  but  of  almost  uniform  number , 
in  each.  In  germination  these  are  brought  out  of 
the  soil  by  the  elongation  of  the  caulicle,  and  when 
the  husk  of  the  seed  is  thrown  off  they  expand 
into  a  circle  of  needle-shaped  leaves.  In  the  Pine 
tribe,  all  the  subsequent  leaves  are  alternate  (spiral)  in  arrange- 
ment, with  some  disguises.  In  the  Cypress  tribe,  the  cotyledons 
are  fewer  (not  more  than  four,  and  more  commonly  only  two), 
and  the  subsequent  leaves  also  are  in  whorls  of  two  to  four ; 
t.  e.,  are  either  opposite  or  verticillate.  From  the  occasional 
union  at  base  of  the  cotyledons  of  a  polycotyledonous  embryo  in 
pairs  or  groups,  and  from  a  study  of  their  early  development, 
Duchartre 1  plausibly  maintains  that  such  cotyledons  really  consist 
of  a  single  pair,  parted  into  divisions  or  lobes.  The  ordinary 
interpretation,  however,  is  equally  tenable. 

39.  The  Monocotyledonous  Embryo,  although  theoretically  the 
simplest,  is  practically  a  more  difficult  study.     It  has  a  single 
cotyledon  (as  the  name  denotes)  ;  also  a  single  leaf  to  each  node 

1  Ann.  Sci.  Nat.  ser.  3,  x.  207.     This  view,  which  originated  with  Jussieu, 
is  adopted  by  Parlatore  in  DC.  Prodr.  xvi. 

FIG.  48.    Section  of  a  seed  of  a  Pine,  with  its  embryo  of  several  cotyledons.  49.  Early 
seedling  Pine,  with  its  stemlet,  displaying  its  six  seed-leaves. 


24 


MORPHOLOGY 


of  the  plumule  ;  that  is,  the  leaves  of  the  embryo  are  alternate. 
But  the  caulicle  is  usually  very  short,  and  there  is  no  external 
mark  by  which  its  limits  may  be  distin- 
guished from  the  cotyledon,  until  germi- 
nation has  begun.  For  a  type  of  it,  the 
embryo  of  some  aquatic  or  marsh  plants 
\i  \  //  ffSjffl  may  be  taken,  where  it  forms  the  whole 
11  kernel  of  the  seed  (Fig.  50-53),  and 
80  n.  52  us  the  structure  can  be  made  out  antecedent 
to  germination.  It  is  understood  by  supposing  that  the  cotyle- 
don, which  forms  its  principal  bulk  (the  caulicle  being  only  the 
very  short  thickish  base) ,  is  convolute  around  a  short 
plumule,  and  the  margins  concreted,  except  a  minute 
longitudinal  chink  at  base,  out  of  which  the  growing 
plumule  protrudes  in  germination.  The  embryo  of 
Iris  may  be  similar  in  structure,  but  no  distinction 
of  parts  is  visible.  It  is  very  small  in  proportion  to 
the  size  of  the  seed,  the  kernel  being  mostly  albu- 
men, —  a  supply  of  food,  from  which  the  germinating 
embryo  draws  the  materials  of  its  growth.  When 
this  takes  place,  either  the  cotyledon  or  the  whole 
embryo  lengthens,  its  lower  part  is  pushed  out  of  the 
seed,  a  root  forms  at  the  free  end  of  the  excessively 
short  caulicle,  and  the  plumule  develops  from  the 
other  in  a  series  of  one-leaved  nodes,  the  internodes 
of  which  remain  so  short  that  the  leaves  continue 
in  close  contact,  the  bases  of  the  older  successively 
enclosing  the  inner  and  younger.  (Fig.  55.)  Here, 
therefore,  the  cotyledon  mainly  remains  in  the  seed, 
and  the  seed  remains  underground  (hypogaeous) . 

40.  It  is  somewhat  different  in  the  Onion,  which 
has  a  similar  embryo,  except  that  it  is  longer,  and 
the  cotyledon  is  curved  in  the  albumen  of  the  seed. 
The  first  steps  are  the  same  as  in  Iris  ;  but  as  soon 
as  a  root  is  formed  and  embedded  in  the  soil,  the 
cotyledon  lengthens  vastly  more,  into  a  long  and 
filiform  green  leaf,  which,  taking  an  erect  position, 

FIG.  50.  Seed  of  Triglochin  palustre ;  the  rhaphe,  leading  to  the  strong  chalaza  at  the 
summit,  turned  towards  the  eye.  51.  The  embryo  detached  from  the  seed-coats,  showing 
the  longitudinal  chink  at  the  base  of  the  cotyledon ;  the  short  part  below  is  the  radicle. 
62.  Same,  with  the  chink  turned  laterally,  and  half  the  cotyledon  cut  away,  bringing  to 
Tiew  the  plumule  concealed  within.  53.  A  cross-section  through  the  plumule,  more 
magnified. 

FIG.  54.  Section  of  seed  of  Iris,  enlarged,  showing  the  small  and  apparently  simple 
embryo  at  the  base  of  the  albumen.  55.  Germinating  seed  and  seedling  of  the  same,  of 
natural  size. 


OF  THE   EMBRYO   AND   SEEDLING. 


25 


carries  up  the  light  seed  far  above  the  surface  of  the  ground,  the 
tip  only  remaining  in  the  albumen  of  the  seed  until  that  is  ex- 
hausted, when  the  tip  perishes  and  the  emptied  husk  falls  away. 
About  this  time  the  plumule  shoots  forth  from  one  side  of  the 
subterranean  base  of  this  cotyledonar  leaf,  in  the  form  of  a  second 
and  similar  filiform  leaf,  to  be  followed  by  a  third,  and  so  on. 
The  sheathing  bases  of  these  succeeding  leaves  become  the  coats 
of  the  Onion-bulb.  The  internodes  remain  undeveloped  until  the 
plant  is  ready  to  blossom.  Very  similar  is  the  germination  of 
a  date-seed,  except  that  the 
protruding  cotyledon  does 
not  lengthen  so  much,  nor 
does  it  elevate  the  heavy 
seed.  Instead  of  the  seed 
being  carried  up,  the  lower 
end  of  the  embryo,  contain- 
ing the  plumule,  is  pushed  down  more  or  less  into  the  loose 
soil,  from  which  in  time  the  developing  plumule  emerges. 

41.  The  embryo  of  Grasses,  especially  of  those  which  yield 
the  cereal  grains,  is  more  complex,  owing  mainly  to  the  great  de- 
velopment of  the  plumule 
and  the  manner  in  which  e- 
its  rudimentary  leaves 
successively  enclose  each 
other.  That  of  Maize  or 
Indian  Corn,  one  of  the 
largest,  is  most  convenient 
for  study.  (Fig.  56-59.)  The  floury  part  of  the  seed,  which 
makes  most  of  its  bulk,  is  the  albumen,  largely  composed  of 
starch.  The  embryo  is  exterior  to  this,  applied  to  one  of  its 
flat  sides,  and  reaching  from  the  thinner  edge  to  or  above  the 
middle  in  the  common  variety  of  corn  here  represented.  The 
form  of  the  embryo  is  best  shown,  detached  entire,  in  Fig.  58  : 
its  structure  appears  in  the  sections.  The  outer  part  is  the 
cotyledon,  which  incompletely  enwraps  the  plumule  :  it  adheres 
closely  to  the  albumen  by  the  whole  back,  and  remains  un- 
changed in  germination  :  its  function  is  to  absorb  nutritive 


FIG.  56.  Section,  flatwise,  of  a  grain  of  Indian  Corn,  dividing  the  albumen  and  the 
embryo.  57.  Similar  section  at  right  angles  to  the  first.  58.  A  detached  embryo : 
corresponding  parts  of  Fig.  57  and  58  indicated  by  dotted  lines. 

FIG.  59.  Vertical  section  of  Indian  Corn  across  the  thickness  of  the  grain,  dividing 
the  embryo  through  the  centre  and  displaying  its  parts  :  c,  cotyledon  ;  p.  plumule  ; 
r,  the  radicle  or  caulicle. 

FIG.  60.  Similar  section  of  grain  of  rice.  61.  Same  of  an  oat-grain ;  the  parts 
as  in  Fig.  59. 


26 


MORPHOLOGY 


matter  furnished   by  the  albumen,  and  to  transmit  it  to  the 
growing  plumule.     The  plumule   consists   of  a  succession   of 

rudimentary  leaves,  sheathing 
and  enclosing  one  another,  on 
the  summit  of  a  very  short 
axis,  which  is  mainly  the 
caulicle,  otherwise  called  rad- 
icle. This  is  completely  en- 
closed by  a  basal  portion  of 
the  cotyledon  and  of  the 
outermost  leaf  of  the  plu- 
mule, which  form  a  peculiar 
sheath  for  it,  named  the 
Coleorhiza?  i.  e.  root-sheath  : 
consequently  the  first  root  or 
roots  have  to  break  through 
this  covering.*  As  in  the  Oak 
and  Pea(32),  the  very  first  or 
outermost  leaves  of  the  plu- 
mule develop  Imperfectly  and  not  into 
efficient  foliage.  The  one  in  Fig.  62, 
which  encloses  the  rest  in  the  early 
growth,  is  left  behind  as  a  mere  sheath 
to  the  base  of  the  following  and  more 
perfect  leaves :  it  is  the  same  as  the 
lowest  in  Fig.  63.  The  leaves  are  first 
developed  :  the  internodes  lengthen  later,  and  the  lowest  lengthen 
very  little.  Not  rarely  the  first  root  starts  singly  from  the  tip  of 
the  caulicle  (Fig.  62,  just  as  in  Fig.  55)  ;  but  others  of  equal 
strength  follow  from  any  part  of  the  caulicle,  and  soon  from 
the  nodes  above ;  and  no  tap-root  is  ever  formed. 

42.  A  Pseudo-monocotyledonous  embryo  occasionally  occurs  ; 
that  is,  one  of  the  dicotyledonous  type,  of  which  one  cotyledon  is 
wanting  through  abortion.  This  occurs  in  Abronia,  a  genus 
related  to  Mirabilis,  and  bearing  an  embryo  very  similar  to  that 
represented  in  Fig.  17,  18,  except  that  one  cotyledon  is  absent. 
The  anomaly  of  an  acotyledonous  embryo  occurs  in  Dodder,  a 
plant  of  the  dicotyledonous  type,  but  with  both  cotyledons 

1  This,  the  Coleorhize  of  Mirbel,  should  not  be  confounded  (as  by  some  it 
has  been)  with  the  "root-cap,"  or  tissue  which  ordinary  roots  (whether 
primary  or  secondary)  break  through  in  their  development  or  carry  on 
their  apex. 

FIG.  62.  Early  germination  of  Indian  Corn.  63.  More  advanced  germination  of 
same :  roots  produced  from  portion  of  stem  above  the  cotyledon  as  well  as  below. 


OF  THE  ORGANS  OF  VEGETATION.         27 

actually  wanting,  —  a  correlation  with  its  parasitic  mode  of  life. 
(64,  Fig.  78.) 

43.  The  dicotyledonous  and  the  monocotyledonous  character 
of  the  embryo  is  correlated  with  profound  differences  in  the  whole 
ulterior  development,  as  revealed  in  the  structure  of  the  stem, 
leaves,  and  flower ;  which  differences  mark  the  two  great  divisions 
of  Phaenogamous  plants,  viz.  DICOTYLEDONES  or  DICOTYLEDONOUS 
PLANTS,  and  MONOCOTYLEDONES  or  MONOCOTYLEDONOUS  PLANTS, 
—  names  introduced  into  classification  by  Ray,  and  adopted  by 
A.  L.  Jussieu,  in  his  Genera  Plantarum. 


CHAPTER  III. 

MORPHOLOGY  AND   STRUCTURE   OF  THE   ORGANS   OF  THE 
PLANT  IN  VEGETATION. 

SECTION  I.     OF  THE  ROOT. 

44.  The  Root,  which  has  been  called  the  descending  axis,  is 
that  portion  of  the  body  of  the  plant  which  grows  downward, 
ordinarily  fixing  the  vegetable  to  the  soil,  and  absorbing  from  it 
materials  which  the  plant  may  elaborate  into  nourishment.  As 
already  stated  (24),  the  root  grows  in  length  by  continuous 
additions  of  new  fabric  to  its  lower  extremity,  elongating  from 
that  part  only  or  chiefly ;  so  that  the  tip  of 
a  growing  root  always  consists  of  the  most 
newly  formed  and  active  tissue.  It  normally 
begins,  in  germination,  at  the  root-end  of  the 
caulicle,  or  so  called  radicle.  But  roots  soon 
proceed,  or  may  proceed,  from  other  parts  of 
the  stem,  when  this  is  favorably  situated  for 
their  production.  The  root  does  not  grow 
from  its  naked  apex,  but  from  a  stratum 
immediately  behind  it :  consequently  its  blunt 
or  obtusely  conical  advancing  tip  consists  of  older,  firmer,  and 
in  part  effete  tissue.  The  tip  of  all  secondary  roots  and  rootlets 

FIG.  64.  Magnified  tip  of  root  of  a  seedling  Maple  (such  as  in  Fig.  9),  sufficiently 
enlarged  to  indicate  the  cellular  structure:  a.  the  portion  where  growth  la  taking 
place;  b.  the  older  and  firmer  tip. 


28 


MORPHOLOGY   OF  THE  ROOT. 


is  similarly  capped  or  protected.1  But  the  so-called  root-cap  is 
seldom  so  distinct  or  separable  as  to  deserve  a  particular  name. 
45.  Nature  of  Growth,  Cells.  The  development  and  growth  of 
the  root,  as  of  other  organs,  results  from  the  development, 
growth,  and  increase  in  number  of  certain  minute  parts,  of  which 
the  plant  is  built  up.  These  component  parts  are  so  much  alike, 
at  least  in  an  early  stage,  and  are  so  obviously  formed  all  on 
one  type,  that  they  take  one  common  name,  that  of  CELLS. 
These  are  the  histological  elements  of  plants,  i.  e.  the  units  of 
minute  anatomical  structure.  While,  in  the  morphology  of  the 
plant's  obvious  organs,  analysis  brings  us  to  the  phytomer  (16) 
as  the  individual  element  which  by  a  kind  of  propagation 
produces  its  like  in  a  second  phytomer,  remaining  however  in 
connection  with  the  first,  thus  building  up  the  general  structure, 
so,  in  an  analogous  way,  each  of  the  obvious  parts  —  each  stalk 
or  blade  or  rootlet  —  is  microscopically  determined  to  be  com- 
posed of  these  ultimate  organic  units,  generally  called  cells. 
The  cell  (cellula,  by  the  French 
conveniently  termed  cellule)  is  the 
living  vegetable  unit,  in  the  same 
sense  that  the  brick  is  the  unit 
of  a  brick  edifice.  To  make  this 
analogy  fairly  complete,  the 
bricks  should  be  imagined  to 
have  a  firm  exterior  or  shell, 
and  a  soft  or  at  length  hollow 
_  interior,  also  to  be  living  when 
incorporated  into  the  structure, 
and  finally  to  be  produced  in  the 
forming  structure  by  a  kind  of 
propagation.  The  production  or 
increase  in  number  of  these  cells  by  development  from  previous 
ones,  and  their  successive  increase  in  size  up  to  maturity,  are 
what  constitutes  vegetable  growth.2  The  inspection  through  a 

1  The  notion  that  the  tip  of  the  root  consists  of  delicate  forming  or 
newly  formed  tissue,  t>r  bears  some  organ  or  structure  of  this  nature  (a 
"  Spongiole  "),  has  hardly  yet  been  eliminated  from  the  text-books  and  popular 
writings.  It  had  no  proper  foundation  in  fact. 

In  Lemna,  and  in  some  other  aquatics,  and  also  in  some  aerial  roots,  this 
older  tissue  often  separates  into  a  real  root-cap,  free  at  base,  like  an  inverted 
calyptra. 

3  This,  as  to  the  structure,  is  the  subject  of  Histology ;  as  to  processes  or 
actions, the  subject  of  Physiology;  both  to  be  treated  in  a  separate  volume. 

FIG.  65,  66.  Portions  of  surface  of  Fig.  64,  more  magnified,  clearly  displaying  the 
superficial  cellular  structure  and  the  long  processes  from  some  of  the  cells,  called  root- 
hairs,  which  abound  on  the  upper  part  of  Fig.  64 


MORPHOLOGY   OF  THE  ROOT.  29 

simple  microscope  of  a  slender  young  root,  and  of  thin  slices 
of  it  immersed  in  water,  may  serve  to  give  a  general  though 
crude  idea  of  the  vegetable  cellular  structure,  sufficient  for  the 
present  purpose.  Roots  are  naked  ;  that  is,  they  bear  no  other 
organs.  When  they  send  off  branches,  these  originate  from  the 
main  root  just  as  roots  originate  from  the  stem ;  and  in  both 
cases  without  much  predetermined  order.  The  ultimate  and 
very  slender  branches  are  sometimes  called  root-fibrils ;  but 
these  are  only  delicate  ramifications  of  the  root.  Like  any 
other  part  of  the  plant,  however,  roots  may  produce  hairs  or 
such  like  growths  from  the  surface,  which  are  wholly  distinct 
from  branches.  (383.) 

46.  Root- hairs.     Roots  absorb  water,  &c.,  from  the  soil  by 
imbibition  through  the  surface  ;  that  is,  through  the  walls  of  the 
eells,  which  are  in  a  certain  sense  permeable  to  fluids,  more  readily 
when  young  and  tender,  less  so  when  older  and  firmer.     Roots, 
therefore,  absorb  most  by  their  fresh  tips  and  adjacent  parts ; 
and  these  are  continually  renewed  in  growth  and  extended  fur- 
ther into  the  soil.     As  the  active  surface  of  a  plant  above  ground 
is  enormously  increased  by  the  spread  of  foliage,  so  in  a  less 
degree  is  the  absorbing  surface  of  young  roots  increased  by  the 
production  of  root-hairs.  (Fig.  64,  upper  part,  and  more  magni- 
fied in  Fig.  65,  66.)     These  are  attenuated  outgrowths  of  some 
part  of  the  superficial  cells  into  capillary  tubes  (only  one  from 
each  cell) ,  closed  at  the  tip,  but  the  calibre  at  base  continuous 
with  the  cavity  of  the  cell ;  into  which,  therefore,  whatever  is 
imbibed  through  the  thin  wall  may  freely  pass.     These  appear 
(as  Fig.  64  shows)   at  a  certain  distance  behind  the  root-tip. 
Further  back  the  older  or  effete  root-hairs  die  away  as  the  cells 
which  bear  them  thicken  into  a  firmer  epidermis. 

47.  To  the  general  statement  that  roots  give  birth  to  no  other 
organs,  there  is  this  abnormal,  but  by  no  means  unusual  excep- 
tion, that  of  producing  buds,  and  therefore  of  sending  up  leafy 
branches.     Although  not  naturally  furnished  with  buds  in  the 
manner  of  the  stem,  yet  many  roots  have  the  power  of  originat- 
ing them  under  certain  circumstances,  and  some  produce  them 
habitually.     Thus  Apple-trees  and  Poplars  send  up  shoots  from 
the  ground,  especially  when  the  superficial  roots  are  wounded. 
And  the  roots  of  Maclura  or  Osage  Orange  so  readily  originate 
buds  that  the  tree  is  commonly  propagated  by  root-cuttings. 

48.  Kinds  of  Roots.     The  root,  commonly  single,  which  origi- 
nates from  the  embryo  itself,  is  called  the  PRIMARY  ROOT.  (37.) 
Roots  which  originate  from  other  and  later  parts  of  the  stem, 
or  elsewhere,  are  distinguished  as  SECONDARY  ROOTS.     But  the 


30  MORPHOLOGY  OP   THE  BOOT. 

latter  are  as  normal  as  the  primary  root ;  that  is,  to  stems 
so  situated  that  they  can  produce  them.  Most  creeping  plants 
emit  them  freely,  usually  from  the  nodes ;  and  so  do  most 
branches,  not  too  old,  when  bent  to  the  ground  and  covered 
with  earth,  thus  securing  the  requisite  moisture  and  darkness. 
Separate  pieces  of  young  stems  (cuttings)  can  commonly  be 
made  to  strike  root.  Upon  this  faculty  of  stems  to  originate 
roots  depends  all  propagation  by  division,  by  laying  or  layering, 
by  cuttings,  &c.  It  is  mainly  annuals  and  common  trees  that 
naturally  depend  on  the  primary  root ;  and  most  of  these  can  be 
made  to  produce  secondary  roots.  Even  leaves  and  leaf-stalks 
of  some  plants  may  be  made  to  strike  root  and  be  used  as 
cuttings.  (77.) 

49.  Duration.     By  differences  in  respect  to  this,  either  the 
root  or  the  plant,  as  the  case  may  be,  is  distinguished   into 
Annual,  Biennial,  or  Perennial,  according  to  whether  life  is  contin- 
ued for  a  single  year  or  season,  for  two,  or  for  a  greater  number. 
The  difference  is  not  in  all  cases  absolute  or  even  well  marked. 

50.  Annuals  are  plants  which,  springing  from  the  seed,  flower 
and  seed  the  same  year  or  season,  and  die  at  or  before  its  close. 
They  produce  fibrous  roots,  either  directly  from  the  embryo  and 
succeeding  joints  of  stem  (as  in  Grasses,  Fig.  63),  or  from  a 
persistent  primary  or  tap-root,  more  or  less  thickened  into  a  trunk 
or  divided  into  branches.     The  products  of  vegetation  in  all  such 
herbs  are  not  stored  in  subterranean  or  other  reservoirs,  but  are 
expended  directly  in  new  vegetative  growth,  in  the  production 
of  blossom,  and  finally  in  the  maturation  of  fruit  and  seed. 
This  completed,  the  exhausted  and  not  at  all  replenished  indi- 
vidual perishes. 

51.  But  some  annuals  may  have  their  existence  prolonged  by 
not  allowing  them  to  blossom  or  seed.     Others,  with  prostrate 
stem  or  branches,  may  from  these   produce  secondar}7  roots, 
which,  forming  new  connections  with  the  soil,  enable  the  newer 
growth  to  survive  when  the  older  parts  with  the  original  root 
have  perished.   And  many  herbs,  naturally  annuals,  are  continued 
from  year  to  }rear  through  such  propagation  from  the  branches, 
used  as  layers  or  cuttings.     Moreover,  certain  plants  (such  as 
Ricinus  or  Castor-oil  Plant) ,  which  are  perennial  or  even  arbo- 
rescent in  warm  climates  to  which  they  belong,  become  annuals 
in  temperate  climates,  early  perishing  by  autumnal  cold. 

52.  The  annuals  of  cool  climates,  where  growth  completely 
ceases  in  winter,  germinate  in  spring,  mature,  and  die  in  or  before 
autumn.     But,  in  climates  with  comparatively  warm  and  rainy 
winter  and  rainless  summer,  many  germinate  in  autumn,  vegetate 


MORPHOLOGY  OF  THE  BOOT.  31 

through  the  winter,   flower  and  seed  in  spring,  and  perish  in 
early  summer.     These  may  be  termed  WINTER  ANNUALS. 

53.  Biennials  are  plants  which,  springing  from  the  seed  and 
vegetating  in  one  season,  live  through  the  interruption  of  winter, 
and  blossom,  fructify,  and  perish  in  the  next  growing  season ; 
their  life  being  thus  divided  into  two  stages,  the  first  of  vegeta- 
tion, the  second  of  fructification.  In  typical  biennials,  nearly 
the  whole  work  of  vegetation  is  accomplished  in  the  first  stage, 
with  the  result  of  accumulation  of  a  stock  of  nutritive  matter, 
to  be  expended  in  the  second  stage  in  the  production  of  blossom 
and  seed.  This  accumulation  is  usually  stored  in  the  root  or  in 
the  base  of  a  very  short  stem  in  connection  with  the  root.  The 
root  of  a  biennial  accordingly  enlarges  and  becomes  fleshy,  or 
obese,  as  this  matter  accumulates.  At  the  close  of  the  growing 
season,  —  the  leaves  perishing  and  the 
stem  having  remained  very  short  (with 
undeveloped  internodes) ,  —  the  root, 
crowned  with  the  bud  or  buds,  contains 
the  main  result  of  the  summer's  work, 
as  provision  for  the  next  year's  devel- 
opment and  the  completion  of  the 
cycle.  This  development,  being  thus 
amply  provided  for,  is  undertaken  in 
spring  with  great  vigor ;  blossom,  fruit, 
and  seed  are  rapidly  produced ;  and 
the  stock  being  consumed,  but  not  at 
all  replenished,  the  cells  of  the  great 
root  are  now  empty  and  effete,  and 
the  individual  perishes.  The  Beet, 
Turnip,  Parsnip,  and  Carrot  are  fa- 
miliar examples  of  biennials,  with  the 
store  of  nourishment  in  the  root.1 
The  Kohl-rabi  is  a  biennial  with  this  deposit  in  the  stem: 
the  Cabbage,  partly  in  the  stem,  partly  in  the  head  of  leaves. 

1  In  these  the  caulicle  enlarges  with  the  root,  so  that  the  upper  and 
bud-bearing  end  is  stem. 

Tap-roots  of  this  kind  are  said,  in  descriptire  botany,  to  be 

Fusiform  or  Spindle-shaped,  when  broader  in  the  middle  and  tapering 
towards  both  ends,  as  in  the  common  Radish  (Fig.  67); 

Conical,  when  tapering  regularly  from  base  to  tip,  as  in  carrots,  &c. ; 

Napiform,  i.  e.  Turnip-shaped,  when  the  thickened  part  is  wider  than  high,  &c. 

Fascicled  Roots  are  those  which  form  in  clusters ;  these  may  be  slender  or 
thickened.  When  much  thickened,  either  irregularly  or  not  of  the  abore 
shapes,  they  are  said  to  be  tuberous. 

FIG.  67.    Radish:  a  fusiform  tap-root 


32  MORPHOLOGY  OF  THE  ROOT. 

54.  But  some  plants,  such  as  the  Radish,  which  when  they 
spring  from  seed  in  autumn  are  true  biennials,  will  when  raised 
in  spring  pass  on  directly  to  the  flowering  stage  in  summer,  or 
when  sown  after  the  warm  season  begins  will  often  run  through 
their  course  as  annuals.     Then  there  are  various  biennials  which 
thicken  the  root  very  little  and  hold  their  leaves  through  the 
winter.     Between  these  and  winter  annuals  no  clear  demarcation 
can  be  drawn.     As  respects  annual  and  biennial  duration,  the 
terms  may  for  the  most  part  be  applied  indiscriminately  to  the 
plant  or  to  the  root.     We  may  say  either  that  the  plant  is  a 
biennial,  or  that  its  root  is  biennial. 

55.  Perennials  are  plants  which  live  and  blossom  or  fructify 
year  after  year.     They  may  or  they  may  not  have  perennial 

roots.  In  trees  and  shrubs,  also  in 
herbs  with  growth  from  year  to  year 
from  a  strong  tap-root,  the  root 
is  naturally  perennial.  But  in  most 
perennials  with  only  fibrous  roots, 
these  are  produced  anew  from  time 
to  time  or  from  year  to  year.  Also, 
while  some  such  roots  remain  fibrous 
and  serve  only  for  absorption,  others 
ma}'  thicken  in  the  manner  of  the 
ordinary  biennial  root  and  serve  a 
similar  use,  i.  e.  become  reservoirs  of 
elaborated  nourishment.  The  Dahlia 
(Fig.  68)  and  the  Peony  afford  good 
examples  of  this.  Sweet  potato  is 
another  instance.1  IVlost  such  roots 
have  only  a  biennial  duration :  they 
are  produced  in  one  growing  season ;  they  yield  their  store  to 
form  or  aid  the  growth  of  the  next.  When  perennials  store  up 
nutritive  matter  underground,  the  deposit  is  more  commonly 
made  in  a  subterranean  portion  of  the  stem,  in  tubers,  corms, 
bulbs,  &c.  (See  115-122.) 

56.  The  distinction  between  annuals  and  biennials  is  at  times 
so  difficult,  and  the  particular  in  which  they  agree  so  manifest, 
—  namely,  that  of  blossoming  only  once,  then  dying,  as  it  were 
by  exhaustion, — that  it  was  proposed  by  DeCandolle  to  unite 

1  It  is  only  by  the  readiness  of  this  root  to  produce  adventitious  buds, 
especially  from  its  upper  part,  that  it  has  been  mistaken  for  a  tuber,  such 
as  the  common  potato. 

FIG.  68.  Fascicled  and  tuberous  or  fusiform  (secondary)  roots  of  Dahlia :  a,  a.  bud* 
on  ba»e  of  the  stem. 


MORPHOLOGY   OF  THE   BOOT.  33 

the  two  under  the  common  appellation  of  MONOCAEPIC  plants, 
Plant fe  monocarpicce ,  taken  in  the  sense  of  only  once-fruiting 
plants ;  and  to  designate  perennials  by  the  corresponding  term 
of  POLYCARPIC,  Plantce  polycarpicas,  literally  many-fruited,  taken 
in  the  sense  of  many-times  fruiting.1 

57.  But  the  distinction  even  here  is  no  more  absolute  than 
that  between  annuals  and  biennials.      For  example,  it  is  not 
quite  clear  whether  the  Cardinal  Flower  and  related  species  of 
Lobelia  should  be  ranked  as  annuals,  biennials,  or  perennials. 
The  plants  may  blossom  and  seed  toward  the  end  of  the  season 
in  which  they  carne  from  seed ;  or,  germinated  in  autumn,  the 
small  seedlings  may  survive  the  winter ;  but  whenever  fructified 
the  fibrous-rooted  mother  plant  dies  throughout ;  yet  usually  not 
before  it  has  established,  and  perhaps  detached  from  the  base, 
small  offsets  to  blossom  the  next  season ;    and  so  on.     Then 
Houseleeks  (Sempervivum)  and  such-like  fibrous-rooted  succu- 
lent plants  multiply  freely  by  offsets  which  are  truly  perennial 
in  the  sense  that  they  live  and  grow  for  a  few  or  several  years  ; 
but  when  at  length  a  flowering  stem  is  sent  up  producing  blos- 
som and  seed,  that  plant  dies  as  completely  and  in  the  same 
manner  as  any  biennial,  only  the  generation  of  offsets  surviving. 
The  same  is  true  of  the   Century   plant   (Agave  Americana, 
wrongly  denominated  American  Aloe),  which  vegetates  in  the 
manner  of  the  accumulating  stage  of  a  biennial,  except  that 
this  continues  for  several  or  very  many  years,  while  the  flower- 
ing stage,  when  it  arrives,  is  precipitated  and  terminated  in  a 
single  season. 

58.  Although  the  stem  usualty  sends  forth  roots  only  when 
covered  by  or  resting  on  the  soil,  which  affords  congenial  dark- 
ness and  moisture,  yet  these  are  in  some  cases  produced  in  the 
open  air.     Roots  may  likewise  subserve  other  and  more  special 
uses  than  the  absorption  of  crude  or  the  storing  of  elaborated 
nourishment. 

59.  Aerial  Roots  is  a  general  name  for  those  which  are  pro- 
duced in  the  open  air.     One  class  of  these  may  serve  the  office 
of  ordinary  roots,  by  descending  to  the  ground  and  becoming 
established  in  the  soil.     This  occurs,  on  a  small  scale,  in  the 
stems  of  Indian  Corn ;  the  lower  nodes  emitting  roots  which 
grow  to  the  length  of  several  inches  before  they  reach  the  ground 


1  These  terms  or  some  equivalents  have  a  convenience  in  descriptive 
botany.     But  those  employed  by  DeCandolle  are  not  happily  chosen,  as  has 
often  been  said.     Mondtocous  (bearing  progeny  once)  and  Pdytocotis  (bearing 
many  times)  would  be  more  appropriate. 
8 


34 


MOEPHOLOGY  OF   THE   ROOT. 


into  which  they  penetrate.  More  remarkable  cases  abound  in 
those  tropical  regions  where  the  sultry  air,  saturated  with  moist- 
ure for  a  large  part  of  the  year,  favors  the  utmost  luxuriance  of 
vegetation.  In  the  Palm-like  Pandanus  or  Screw-Pine *  (Fig. 
69),  very  strong  roots,  emitted  in  the  open  air  from  the  trunk, 

and  soon  reaching 
the  soil,  give  the 
appearance  of  a  tree 
partially  raised  out 
of  the  ground.  The 
famous  Banyan-tree 
of  India  (Fig.  71)  is 
a  still  more  striking 
illustration ;  for  the 
aerial  roots  strike 
from  the  horizontal 
branches  of  the  tree, 
often  at  a  great 
height,  at  first  swing- 
ing free  in  the  air, 
but  finally  reach- 
ing and  establishing 
themselves  in  the 
ground,  where  they 
increase  in  diameter 
and  form  accessory 
trunks,  surrounding 
the  original  bole  and  supporting  the  wide-spread  canopy  of 
branches  and  foliage.  Very  similar  is  the  economy  of  the  Man- 
grove (Fig.  70) ,  which  forms  impenetrable  thickets  on  low  and 
muddy  sea-shores  in  the  tropics  throughout  most  parts  of  the 
world,  extending  even  to  the  coast  of  Florida  and  Louisiana. 
Here  aerial  roots  spring  not  only  from  the  main  trunk,  as  in 
the  Pandanus,  but  also  from  the  branchlets,  as  in  the  Banyan. 
Even  the  radicle  of  the  embryo  starts  into  growth,  protrudes, 
and  attains  considerable  length  while  the  fruit  is  still  attached  to 
the  branch. 

59".  Aerial  Rootlets  for  climbing  are  familiar  in  the  Ivy  of  the 
Old  World  (Hedera) ,  Trumpet-Creeper  (Tecoma  radicans) ,  and 
our  Poison  Ivy  (Rhus  Toxicodendron)  ;  by  the  adhesion  of 

1  So  named,  not  from  any  resemblance  to  a  Pine-tree,  but  from  a  like- 
ness of  the  foliage  to  that  of  a  Pine-Apple. 

FIG.  69.  Pandanus,  or  Screw-Pine;  and  in  the  background,  70,  a  Mangrove-tree 
(Rhizophora  Mangle). 


MORPHOLOGY  OF  THE   BOOT.  35 

which  the  stems,  as  they  grow,  ascend  walls  and  the  trunks  of 
trees  with  facility.  In  Rhus  a  superabundance  of  these  rootlets 
is  produced,  thickly  covering  ah1  sides  of  the  stem. 


60.  Epiphytes  or  Air-Plants  also  have  roots  which  are  through- 
out life  unconnected  with  the  ground.  Epiphytes,  or  Epiphytic 
plants,  as  the  name  denotes,  are  such  as  grow  upon  other  plants 
without  taking  nourishment  from  them.  Deriving  this  from  the 
air  alone,  they  are  called  Air-plants.  This  name  might  be 
extended  to  the  same  or  other  kinds  of  plants  attaching  them- 
selves to  bare  walls,  rocks,  and  the  like,  and  unconnected  with 
the  soil,  though  such  would  not  technically  be  epiphytes.  Very 
many  Lichens,  Mosses,  and  other  plants  of  the  lower  grade,  and 
not  a  few  phaenogamous  plants,  are  in  this  case.  The  greater 
part  of  the  phaenogamous  Epiphytes  pertain  to  two  monocotyle- 
donous  orders,  the  Orchis  family  and  that  to  which  the  Pine- 
Apple  belongs,  viz.  the  Bromeliaceae.  Their  thread-like  or 
cord-like  simple  roots  either  adhere  to  the  bark  of  the  supporting 
tree,  securing  the  plant  in  its  position,  or  some  hang  loose  in  the 
air.  Of  these,  Orchids,  i.  e.  plants  of  the  Orchis  family,  are  the 
most  showy  and  numerous,  and  of  the  greatest  variety  of  forms, 
especially  of  their  blossoms,  which  are  often  bizarre  and  fantas- 
tic. They  belong,  naturally,  to  climates  which  are  both  warm 
and  humid ;  they  are  highly  prized  in  hot-house  cultivation ; 
arid,  along  with  the  hardy  and  terrestrial  portion  of  the  order, 
they  are  peculiarly  interesting  to  the  botanist  on  account  of  the 
singular  and  exquisite  adaptation  of  their  flowers  in  relation 
to  insects  which  visit  them.  In  some  the  blossoms  curiously 

FIG.  71.    The  Banyan- tree,  or  Indian  Fig  (Ficus  Indica). 


36  MORPHOLOGY  OF   THE  BOOT. 

resemble  butterflies  or  other  insects  ;  as,  for  example,  Oncidium 
Papilio,  Fig.  72.  Epiphytic  orchids  are  indigenous  to  the  United 
States  only  from  Georgia  to  Texas,  and  only  in  humble  forms, 
in  company  with  species  of  Tillandsia,  representing  Bromeliace- 
ous  epiphytes.  The  commonest  of  the  latter  tribe,  and  of  most 
northern  range,  is  the  T.  usneoides,  the  so-called  Long  Moss, 
which,  pendent  in  long  and  tangled  gray  clusters  or  festoons  from 
the  branches  of  the  Live-Oak  or  Long-leaved  Pine,  gives  such  a 
peculiar  and  sombre  aspect  to  the  forests  of  the  warmer  portions 
of  our  Southern  States. 


61.  Parasitic  Plants  have  the  peculiarity  that  their  roots,  or 
what  answer  to  roots,  not  only  fix  themselves  to  other  plants, 
but  draw  therefrom  their  nourishment,  at  least  in  part.     Among 
cryptogamous   plants   very  many  Fungi  are  parasitic  upon  or 
within  living  plants  or  animals.     But  only  phaenogamous  para- 
sites are  here  under  consideration.     These  may  be  divided  into 
two  classes  ;  those  with  and  those  without  green  foliage. 

62.  Green  Parasites  may  be  either  wholly  or  partially  parasitic  ; 
that  is,  they  may  draw  all  their  support  from  a  foster  plant,  or 

FIG.  72.    Oncidium  Papilio,  and,  73,  Comparettia  rosea;  two  showy  epiphytes  of  the 
Orchis  family;  showing  the  mode  in  which  these  Air-plants  grow. 


MORPHOLOGY   OF   THE  BOOT. 


37 


they  may  be  likewise  rooted  in  the  soil,  and  receive  from  it 
materials  of  their  food.  Having  green  foliage,  they  are  capable 
of  elaborating  such  food,  whether  taken  directly  from  the  soil  or 
from  the  crude  sap  of  the  foster  plant.  The  Mistletoes  (Viscum 
and  its  allies)  are  the  principal  examples  of  complete  green 
parasitic  plants.  Seeds  dropped  by  birds  on  the  boughs  of  trees 
germinate  there  ;  the  root-end  of  the  caulicle  points  thither  instead 
of  towards  the  earth ;  the  root,  or  what  would  be  such,  pene- 
trates the  bark  and  in- 
corporates itself  with  the 
sap-wood  so  perfectly 
that  the  junction  of  par- 
asite with  foster  trunk 
is  like  that  of  branch 
with  parent  trunk.  The 
parasite  is  probably  fed 
by  both  elaborated  and 
crude  sap,  that  is,  both 
by  what  the  foster  tree 
has  assimilated  and 
what  it  has  merely  taken 
from  the  soil  and  air: 
the  former  it  can  at  once 
incorporate ;  the  latter 
it  has  first  to  assimilate 
in  its  own  green  leaves. 
Sometimes  one  Mistleto 
is  parasitic  upon  an- 
other of  the  same  or  of 
a  different  species. 

63.  Partially  parasitic  plants  (mostly  green)  may  be  either  woody 
and  arborescent  or  herbaceous.    The  species  of  Clusia  in  tropical 


FIG.  74.    Native  epiphytes  of  Georgia,  &c. :  the  erect  one  at  the  right  an  Orchid, 
Epidendrum  conopseum ;  the  hanging  one  Tillandsia  nsneoides,  called  Long  Moss. 

FIG.  75-    Boots  of  Gerardia  flava:  some  of  the  rootlets  attaching  themselv 
sitically  to  the  root  of  a  Blueberry.    (From  a  drawing  by  Mr.  J.  Stauffer.) 


38 


MORPHOLOGY   OP   THE   ROOT. 


America  (called  Cursed  Fig)  are  examples  of  the  former.  They 
form  trees,  send  down  aerial  roots  in  the  manner  of  the  Banyan  ; 
but,  while  some  roots  seek  the  ground,  some  may  attach  them- 
selves to  other  trees  parasitically,  and  draw  from  them  a  portion 
of  their  support.  The  parasitism  of  certain  herbaceous  plants 
with  green  foliage  is  clandestine,  the  connection  being  under- 
ground and  therefore  long  unsuspected.  This  occurs  in  species 
of  Gerardia  (at  least  of  the  section 
Dasystoma)  and  other  plants  of  the 
same  family,  the  uncultivability  of 
which  is  thereby  explained.  Also 
in  Comandra  and  in  their  relatives 
the  Thesiums  of  the  Old  World, 
belonging  to  a  natural  order  (the  Santalaceae)  which  has  much 
affinity  with  the  entirely  parasitic  order  (Loranthacese)  to  which 
the  Mistleto  belongs. 

64.  Pale  or  Colored  Parasites,  such  as  Beech-Drops,  Pine-Sap, 
&c.,  are  those  which  are  destitute  of  green  herbage,  and  are 
usually  of  a  white,  tawny,  or  reddish 
hue ;  in  fact,  of  any  color  except 
green.  These  strike  their  roots,  or 
sucker-shaped  discs,  into  the  bark, 
mostly  that  of  the  root,  of  other 
plants,  and  thence  draw  their  food 
from  the  sap  already  elaborated. 
They  have  accordingly  no  occasion 
for  digestive  organs  of  their  own, 
i.  e.  for  green  foliage.  The  Dodder 
(Fig.  77)  is  a  common  plant  of 
this  kind  which  is  parasitic  above 
ground.  Its  seeds  germinate  in 
the  earth,  but  form  no  proper  root : 
when  the  slender  twining  stem 
reaches  the  surrounding  herbage, 
it  forms  suckers,  which  attach 
surface  of  the  supporting  plant, 
penetrate  its  epidermis,  and  feed  upon  its  juices ;  while  the 
original  root  and  base  of  the  stem  perish,  and  the  plant  has 
no  longer  any  connection  with  the  soil.  Thus  stealing  its  nour- 

FIG.  76.    Section  of  one  of  the  attached  rootlets  of  Gerardia,  showing  the  union. 

FIG.  77.  The  common  Dodder  of  the  Northern  States  (Cuscuta  Gronovii),  of  the 
natural  size,  parasitic  upon  the  stem  of  an  herb :  the  uncoiled  portion  at  the  lower  end 
shows  the  mode  of  its  attachment.  78.  The  coiled  embryo  taken  from  the  seed,  con- 
sisting of  naked  caulicle  and  plumule ;  moderately  magnified.  79.  The  same  in  germi- 
nation, elongating  into  a  thread-like  leafless  stem. 


MORPHOLOGY   OF  THE  BOOT.  39 

ishment  ready  prepared,  it  requires  no  proper  digestive  organs 
of  its  own,  and,  consequently,  does  not  produce  leaves.  This 
economy  is  foreshadowed  in  the  embryo  of  the  Dodder,  which  is 
a  naked  thread  spirally  coiled  in  the  seed  (Fig.  78,  79),  and 
presenting  no  vestige  of  cotyledons  or  seed-leaves.  A  species 
of  Dodder  infests  and  greatly  injures  flax  in  Europe,  and  some- 
times makes  its  appearance  in  our  own  flax- fields,  having  been 
introduced  with  the  imported  seed.  Such  parasites  do  not  live 
upon  all  plants  indiscriminately,  but  only  upon  those  whose 
elaborate  juices  furnish  a  propitious  nourishment.1  Some  of 
them  are  restricted,  or  nearly  so,  to  a  particular  species  ;  others 
show  little  preference,  or  are  found  indifferently  upon  several 
species  of  different  families.  Their  seeds,  in  some  cases,  it  is 
said,  will  germinate  only  when  in  contact  with  the  stem  or  root 
of  the  species  upon  which  they  are  destined  to  live.  Having  no 
need  of  herbage,  such  plants  may  be  reduced  to  a  stalk  bearing 
a  single  flower  or  a  cluster  of  flowers,  or  even  to  a  single  blossom 
developed  from  a  bud  directly  parasitic  on  the  bark  of  the  foster 
plant.  Of  this  kind  are  the  several  species  of  Pilostyles  (para- 
sitic flowers  on  the  shoots  of  Leguminous  plants)  in  Tropical 
America,  one  species  of  which  was  discovered  by  Dr.  Thurber 


near  the  southern  borders  of  New  Mexico.  Its  flowers  are 
small,  only  about  a  quarter  of  an  inch  in  diameter.  The  most 
wonderful  plant  of  this  kind  is  that  vegetable  Titan,  the  Raf- 
flesia  Arnoldi  of  Sumatra  (Fig.  80)  which  grows  upon  the  stem 
of  a  kind  6f  a  Cissus  or  Vitis.  It  is  a  parasitic  flower,  measuring 
nine  feet  in  circumference,  and  weighing  fifteen  pounds !  Its 
color  is  light  orange,  mottled  with  yellowish- white. 


1  Monotropa  or  Indian  Pipe  (and  perhaps  some  related  plants),  although 
probably  parasitic  on  living  roots  in  early  growth,  appears  to  live  afterwards 
in  the  manner  of  the  larger  Fungi,  upon  leaf-mould  and  decaying  herbage. 
Its  mode  of  life  should  be  investigated. 

FIG.  80.  Rafflesia  Arnoldi ;  an  expanded  flower,  and  a  bud,  directly  pararitic  on 
the  stem  of  a  vine:  reduced  to  the  scale  of  half  an  inch  to  a  foot. 


40  MOBPHOLOGY   OF  BUDS. 

SECTION  II.     OP  BUDS. 

65.  Buds  are  the  germs  of  stems :  they  are  axes  with  their 
appendages  in  an  early  state.     LEAF-BUDS  (GEMWLE)  are  those 
devoted  to  vegetation,  and  the  parts,  or  some  of  them,  develop 
as  leaves.     MIXED   BUDS  contain   both   foliage   and   flower  or 
flowers.    FLOWER-BUDS  (ALABASTRA)  are  unexpanded  blossoms. 
These  are  considered  in  another  chapter. 

66.  The  conspicuous  portion  of  an  ordinary  bud,  or  that  which 
first  develops,  usually  consists  of  leaves,  or  scales  the  homologues 
of  leaves  ;  the  axis  itself  being  very  short  and  undeveloped.     If 
this  remains  comparatively  short,  the  leaves  as  developed  are 
crowded  in  a  rosette,  as  in  a  Houseleek  (Fig.  91*),  a  Barberry 
and  the  Larch :   when  the  internodes  lengthen,  the  leaves  are 
interspaced  upon  the  axis. 

67.  The  cotyledons  and  plumule  of  the  embryo  are,  morpho- 
logically, the  first  bud,  on  the  summit  of  the  initial  stem,  the 
caulicle.     This  in  germination  and  subsequent  growth  develops 
into  a  leafy  stem,  in  the  manner  already  described.     Normally 
this  stem  has  the  capacity  of  growing  on  in  this  way  from  the 
apex  or  growing  point,  which  is  always  potentially  a  bud,  the 
apical  or  terminal  bud  (15).      Sometimes  it  is  merely  potential, 
and  there  is  no  external  structure  visible  until  the  new  growth 
begins,  or  the  bud  is  said  to  be  latent. 

68.  But  commonly,  in  plants  that  live  from  year  to  year,  growth 
is  divided  into  seasons  or  stages,  with  intervals  of  repose.     In 
such  cases,  especially  in  trees  and  shrubs,  instead  of  a  continuous 
succession  of  foliage,  the  period  of  interruption  is  apt  to  be 
marked  by  the  production  of  scales  (Bud-scales,  Perulce,  etc.)  or 
dry  teguments,  which  serve  to  protect  the  tender  rudiments  or 
growing  point  within  during  the  season  of  rest.     This  being  the 
winter-season  in  cold  climates,  Linnaeus  gave  to  such  bud-cover- 
ings the  common  name  of  HIBERNACULUM.     From  the  usually 
squamose  (scale-like)  character  of  this  covering,  such  buds  take 
the  name  of 

69.  Scaly  Buds.     Large  and  strong  ones  of  this  kind,  such  as 
those  of  Horsechestnut,  Magnolia,  Hickory,  Lilac,  &c. ,  may  be 
taken  as  the  type  of  bud.     The  scales  serve  to  protect  the  ten- 
der parts  within  against  injury  from  moisture  and  from  sudden 
changes  in  temperature  during  the  dormant  or  earliest  growing 
state.    To  ward  off  moisture  more  effectually,  they  are  sometimes 
coated  with  a  wax}',  resinous,  or  balsamic  exudation,  as  is  con- 
spicuous on  the  scales  of  the  Horsechestnut,  Balsam-Poplar  or 
Balm  of  Gilead,  and  Balsam-Fir.      To  guard  against  sudden 


MORPHOLOGY   OF   BUDS. 


41 


changes  of  temperature,  they  are  often  lined,  or  the  rudimentary 
leaves  within  invested  with  non-conducting  down  or  wool. 

70.  Nature  of  Bud-scales.     That  they  answer  to  leaves  is  made 
manifest  by   a   consideration   of  their 

situation  and  arrangement,  which  are 
the  same  as  of  the  proper  leaves  of 
the  species  ;  and  by  the  gradual  transi- 
tions from  the  former  to  the  latter  in 
many  plants.  In  the  Turions,  or  sub- 
terranean budding  shoots  of  numerous 
perennial  herbs,  and  in  the  unfolding 
buds  of  the  Lilac  and  Sweet  Buckeye 
(.^Esculus  parviflora),  every  gradation 
may  be  traced  between  bud-scales  and 
foliage,  showing  that  no  line  of  distinc- 
tion can  be  drawn  between  them,  but 
that  the  two  are  essentially  of  the  same 
nature,  are  different  modifications  of 
the  same  organ.  In  the  Lilac  they 
may  be  regarded  as  the  blade  of  the 
leaf,  modified  and  depauperate ;  in  the 
Buckeye  (Fig.  233),  and  therefore  in 
Horsechestnut,  as  the  base  of  leaf-stalks ; 
in  Magnolia  (Fig.  81,  82),  in  the 
Tulip-tree,  and  in  the  Beech,  they  are 
evidently  stipules.  They  must  therefore 
be  referred  to  in  the  section  on  the 
morphology  of  leaves.  (227.) 

71.  Naked  Buds,  &c.,  of  shrubs  and 
trees,    even   in    climates    with    severe 
winter,  are  not  unknown,  that  is,  buds 
unprotected  by  special  scales  or  other 
coverings.      For    example,   the    latest 
pair  of  leaves  of  the  season  in  Viburnum 
nudum,  V.  lantanoides   (Hobblebush) , 
and  the  like,  remain  in  a  nascent  state 
over  winter  without  covering,  and  ex- 
pand into  the  first  foliage  in  the  spring. 

Yet  V.  Opulus  (Snowball,   &c.),  another  species  of  the  same 
genus  and  inhabiting  the  same  region,  has  well-formed  scaly 

FIG.  81.  Branch  of  Magnolia  Umbrella,  of  the  natural  size,  crowned  with  the 
terminal  bud ;  and  below  exhibiting  the  large  rounded  leaf-scars,  as  well  as  the  rings 
or  annular  scars  left  by  the  fall  of  the  bud-scales  of  the  previous  season.  82.  A  detached 
scale  from  a  similar  bud ;  its  thickened  axis  is  the  base  of  a  leaf-stalk ;  the  membranous 
sides  consist  of  the  pair  of  stipules  united  with  it. 


42 


MOKPHOLOGY   OF  BUDS. 


leaf-buds.     In  other  hardy  shrubs  and  trees,  the  buds,  equally 

or  almost  destitute  of  scales, 
are  minute,  hidden  in  or  un- 
der the  bark,  or  otherwise 
inconspicuous  until  vernal 
growth  commences.  Phila- 
delphus  and  Taxodium  are  of 
this  kind. 

72.  Snbpetiolar  Buds.  Some 
leaf-buds  are  singularly  cov- 
ered in  their  early  state  and 
through  the  summer,  as  in  the 
Locust  (Robinia) ,  Honey-Lo- 
cust Fig.  96  (where  they  re- 
main very  undeveloped),  in 
Yellow  Wood  (Cladrastis),and 
more  conspicuously  in  the 
Plane-tree  (Platanus,  Fig.  87) : 
here  they  are  all  formed  un- 
der the  base  of  the  protecting 
leaf-stalk,  which  in  Plane-tree 
forms  a  sheath  or  inverted  cup, 
very  like  a  candle-extinguisher, 

fitted  to  and  concealing  the  conical  bud  until  autumn,  when  by 

the  fall  of  the  leaves  these  buds  are  exposed. 


FIG.  83.  Diagram  of  vertical  section  of  a  strong  bud,  such  as  of  Horsechestnut. 
84.  The  axis  of  the  same  developing,  the  elongation  beginning  with  the  lowest  inter- 
node,  soon  followed  by  the  others  In  succession.  85.  A  year's  growth  of  Horsechestnut, 
crowned  with  a  terminal  bud:  a,  scars  left  by  the  bud-scales  of  the  previous  year: 
b,  scars  left  by  the  fallen  leaf-stalks:  c,  axillary  buds. 

FIG.  86.    Branch  and  buds  (all  axillary)  of  the  Lilac. 

FIG.  87.    Leaf-bud  under  the  petiole  of  the  Plane-tree. 


MOBPHOLOGY  OF  BUDS.  43 

73.  Fleshy  Buds.     Bulbs  are  peculiar  buds  of  certain  herba- 
ceous plants,  with  fleshy  scales,  and  often  of  a  more  permanent 
character.     Their  nature  and  economy  may  most  conveniently 
be  illustrated   under   subsequent   sections.     Usually  bulbs  are 
subterranean  or  partly  so.     But  small  bulbs  (Bulblets,  123)  regu- 
larly appear  in  the  axil  of  nearly  all  the  leaves  of  certain  common 
Lilies,  being  obviously  ordinary  axillary  buds,  under  certain 
modifications.     They  become  detached  at  maturity,  fall  to  the 
ground,  produce  roots,  and  grow  as  independent  plants ;  and 
their  fleshy  scales  are  storehouses  of  nourishment  for  the  early 
support  of  this  independent  growth. 

74.  Bad- propagation  is  a  normal  mode  of  reproduction  in  cases 
like  the  above,  the  spontaneously  detached  bulb  lets  or  buds 
establishing   themselves   as    progeny.      In   several   species   of 
Allium  (Onions  and  Leeks),  such  bulblets  usurp  the  place  of 
flower-buds,  making  the  analogy  seem  closer.    Stems  or  branches 
which  habitually  root  in  the  soil,  or  along  its  surface,  equally 
propagate  or  divide  into  new  individuals,  becoming  distinct  by 
the  perishing  of  the  older  connecting  parts,  or  by  breaking  away 
from  them.     Propagation  by  cuttings  is  an  acceleration  or  exten- 
sion of  this  same  natural  operation.     The  cutting  is  a  portion  of 
stem  bearing  one  or  more  buds,  which,  through  the  faculty  of 
the  stem  to  strike  root,  is  made  to  grow  independently.     In 
grafting,  such  a  cutting,  and  in  budding  a  bud  only,  with  a  small 
portion  of  wood  and  bark,  is  transferred  to  the  stem  of  another 
plant  of  the  same  or  of  some  related  species,  and  made  to  grow 
there,  uniting  its  wood  and  bark  with  those  of  the  stock,  and  so 
becoming  a  limb  or  branch,  in  place  of  striking  root  into  the  soil 
and  becoming  a  separate  plant.     The  horticultural  advantage  of 
bud-propagation  is,  that  the  offsets  or  new  individuals  share 
in  all  the  peculiarities  of  the  parent  as  completely  as  if  still 
branches   of  that  tree.     In   propagation  by  seed,  the  special 
peculiarities  or  excellencies  of  individuals  or  varieties  may  not, 
and  in  some  measure  probably  will  not,  be  reproduced. 

75.  Normal  or  Regular  Buds,  as  to  position,  are  either  terminal 
or  axillary,  as  already  stated.  (15.)     They  are  single,  that  is, 
one  bud  normally  occupies  the  apex  of  a  stem  or  branch,  and 
appears,  or  usually  may  appear,  in  the  axil  of  (or  upper  angle 
formed  with  the  stem  by)  any  well-developed  leaf.     In  these 
positions,  buds  are  so  usual,  or  so  capable  of  appearing,  that 
they  are  commonly  regarded  as   potential   when   not   actually 
present.      The  potentiality  may  be  manifested  by  the  actual 
development  of  these  buds  in  shrubs  or  trees  after  the  lapse  of 
years.  (84.)     The  terminal  leaf-bud  is  to  continue  the  axis  it 


44 


MORPHOLOGY   OP  BUDS. 


surmounts  :  axillary  and  any  other  lateral  leaf-buds  are  to  be- 
come branches.  But  even  of  buds  which  actually  appear  a  large 
proportion  do  not  grow.  When  a  terminal  bud  is  formed  (as 
Fig.  81,  85,  91),  this  is  commonly  the  strongest,  or  among  the 
stronger.  But  in  many  cases  it  habitually  or  commonly  fails  to 
appear.  In  the  Elm  (with  leaves  and  therefore  buds  alternate) , 
the  bud  axillary  to  the  last  leaf  of  the  season  takes  its  place. 
In  the  common  Lilac,  a  pair  of  buds,  which  were  in  the  axils  of 
the  uppermost  of  the  (opposite)  leaves,  seem  to  replace  the 
terminal  bud,  which  seldom  develops.  (Fig.  86.)  When  all  the 
regular  buds  make  their  appearance,  and  the  leaves  are  opposite, 
the  stem  will  be  crowned  with  the  terminal  bud,  having  an  axil- 
lary bud  on  each  side  of  it.  (Fig.  88.) 

76.  Accessory  Buds.     These  are,  as  it  were,  multiplications  of 
the  regular  axillary  bud,  giving  rise  to  two,  three,  or  more,  instead 


0  of  one  ;  in  some  cases  situated  one  above  an* 
other  (superposed),  in  others  placed  side  by 
side  (collateral).  In  the  latter  case,  which 
occurs  occasionally  in  the  Hawthorn,  in  cer- 
tain Willows,  in  the  Maples  (Fig.  88),  &c., 
the  axillary  bud  seems  to  divide  into  three, 
or  itself  to  give  rise  to  a  lateral  bud  on  each 
side.  On  some  shoots  of  the  Tartarean 
Honeysuckle  (Fig.  90)  from  three  to  six  buds 
appear  in  each  axil,  one  above  another,  the 
lower  being  successively  the  stronger  and 
earlier  produced ;  and  the  one  immediately 
in  the  axil,  therefore,  grows  in  preference : 
occasional!}'  two  or  more  of  them  grow,  and 
superposed  accessory  branches  result.  It  is 
much  the  same  in  Aristolochia  Sipho,  except 
that  the  uppermost  bud  is  there  strongest. 


FIG.  88.  Branch  of  Red  Maple,  at  the  middle  bearing  triple  axillary  buds,  placed 
side  by  side. 

FIG.  89.  Piece  of  a  branch  of  the  Butternut,  with  accessory  buds  placed  one  above 
another  •.  a,  the  leaf  scar :  b,  proper  axillary  bud :  a,  d,  accessory  buds. 

FIG.  90.  Part  of  a  branch  of  Tartarean  Honeysuckle,  with  crowded  accessory  buds 
superposed  in  the  axil  of  each  leaf. 


MORPHOLOGY   OF   STEMS.  45 

So  it  is  in  the  Butternut  (Fig.  89),  where  the  true  axillary  bud 
is  minute  and  usually  remains  latent,  while  the  accessory  ones 
are  considerably  remote,  and  the  uppermost,  which  is  much  the 
strongest,  is  far  out  of  the  axil :  this  usually  develops,  and  gives 
rise  to  an  extra-axillary  branch. 

77.  Adventitious  Buds  are  such  as  are  abnormal  and  irregular, 
being  produced  without  order  and  from  any  part  of  the  stem,  or 
even  from  roots.  The  latter,  like  the  internodes  of  a  stem, 
although  normally  destitute  of  buds,  do  produce  them  notwith- 
standing in  certain  cases,  especially  when  wounded,  and  in  some 
plants  (such  as  Blackberries)  so  freely  that  gardeners  propagate 
them  by  root- cuttings.  The  stems  share  this  tendency ;  and 
buds  are  apt  to  break  out  on  the  sides  of  trunks,  especially  when 
wounded  or  pollarded,  or  to  spring  from  new  tissues  produced 
on  cut  surfaces,  especially  where  the  bark  and  wood  join.  Even 
leaves  ma}7  develop  adventitious  buds,  and  then  be  used  for 
propagation.  In  Bryophyllum,  such  buds,  followed  by  rootlets, 
are  freely  produced  on  the  margins  of  the  blade  or  of  its  leaflets. 
In  Begonia,  a  leaf,  used  as  a  cutting,  will  root  from  the  base  of 
the  petiole  stuck  in  the  soil,  and  produce  buds  on  the  blade,  at 
the  junction  with  the  petiole,  or  elsewhere. 


SECTION  III.     OF  THE  STEM. 
§  1.   GENERAL  CHARACTERISTICS  AND  GROWTH. 

78.  The  Stem  is  the  ascending  axis,  or  that  portion  of  the 
trunk  which  in  the  embryo  grows  in  an  opposite  direction  from 
the  root,  seeking  the  light,  and  exposing  itself  as  much  as  pos- 
sible to  the  air.  All  phsenogamous  plants  possess  stems.1  In 
those  which,  in  botanical  descriptions,  are  said  to  be  acaulescent, 
or  stemless,  it  is  either  very  short,  or  concealed  beneath  the 
ground.  Although  the  stem  always  takes  an  ascending  direction 
at  the  commencement  of  its  growth,  it  does  not  uniformly  retain 
it ;  but  sometimes  trails  along  the  surface  of  the  ground,  or 
burrows  beneath  it,  sending  up  branches,  flower-stalks,  or  leaves 
into  the  air.  The  common  idea,  that  all  the  subterra'nean  portion 
of  a  plant  belongs  to  the  root,  is  incorrect.  Equally  incorrect  is 
the  common  expression  that  plants  spring  from  the  root.  Roots 
spring  from  the  stem,  not  the  stem  from  the  root.  (21,  24,  37,  44.) 

1  There  are,  however,  reduced  forms  in  which  there  is  no  distinction  of 
axis  and  foliage ;  but  most  of  these  are  clearly  leafless  rather  than  stemless, 
and  not  even  in  Lemna  and  Wolffia  can  the  stem  be  said  to  be  wanting. 


46  MOKPHOLOGY  OP   STEMS. 

79.  "While  the  root  normally  gives  birth  to  no  other  organs, 
but  itself  performs  those  functions  which  pertain  to  the  relations 
of  the  vegetable  with  the  soil,  —  binding  it  to  the  earth  and 
absorbing  nourishing  materials  from  it,  —  the  aerial  functions  of 
vegetation  are  chiefly  carried  on,  not  so  much  by  the  stem  it- 
self as  by  a  distinct  set  of  organs  which  it  bears,  namely,  the 
leaves.     Hence,  the  production  of  leaves  is  one  of  the  charac- 
teristics of  the  stem.     These  are  produced  only  at  certain  definite 
and  symmetrically  arranged  points,  called  nodes.  (13,  23.) 

80.  Development  and  Structure.    In  a  bud  or  undeveloped  stem, 
the  nodes  are  in  contact  or  close  proximity.     In  the  develop- 
ment, growth  in  length  takes  place  in  such  manner  as  to  carry 
these  apart  more  or  less,  according  to  the  degree  of  elongation, 
that  is,  the  internodes  (13)  elongate.    The  order  of  development 
is  from  below  upward,  the  lowest  internode  first  lengthening, 
the  others  in  regular  succession.     Each  completes  its  growth, 
with  more  or  less  rapidity,  although  the  length  attained  varies 
greatly  in  different  stems,  in  different  parts  of  the  same  stem, 
and  under  different  conditions.     Unlike  the  root,  in  which  the 
elongation  of  formed  parts  is  very  soon  finished  and  therefore 
only  the  tip  is  perceptibly  growing,  internodes  go  on  growing 
throughout,   and  several   formed   internodes   may   be   growing 
simultaneously,  thus  producing  elongation  throughout  a  consid- 
erable extent  of  stem  and  with  considerable  rapidity.     But  each 
internode  grows   independently.     Some  parts  of  an  internode 
may  lengthen  faster  or  continue  in  growth  longer  than  others  ; 
this  is  usually  the  upper  portion,  at  least  in  long  internodes  and 
when  every  part  is  equally  exposed  to  light. 

81.  The  development  of  a  stem  from  a  bud  is  wholly  like  that 
from  the  embryo,  and  has  already  been  described  in  Chap.  II. 
It  exhibits  similar  variations  as  to  rapidity  and  vigor,  dependent 
upon  the  constitution  of  the  bud, — which,  like  the  plumule  in 
the  seed  or  seedling,  may  be  either  latent  or  much  developed 
before  growth  begins,  —  also  upon  the  amount  of  nourishment 
provided.     Strong  buds  commonly  have  their  parts,  or  some  of 
them,  ready  formed  in  miniature,  and  a  store  of  elaborated  nour- 
ishment in  the  parent  stem  to  draw  upon.     Those  well-developed 
buds  which  in  many  of  our  shrubs  and  trees  crown  the  apex  or 
occupy  the  axils  of  stem  and  branches  early  in  the  preceding 
summer  (as  in  Magnolia,  Fig.  81,  Horsechestnut,  Fig.  85,  and 
Hickory,  Fig.  91)    often  exhibit  the  whole  plan  and  amount  of 
the  next  year's  growth ;  the  nodes,  the  leaves  they  bear,  and 
sometimes  the  blossoms  being  already  formed,  and  only  requiring 
the  elongation  of  the  internodes  for  their  full  expansion.     As 


CHARACTERISTICS   AND   GROWTH. 


47 


the  bud  is  well  supplied  with  nourishment  in  spring  by  the  stem 

on  which  it  rests,  its  axis  elongates  rapidly ;  and  although  the 

growth  commences  with  the  lowest  internode,  yet 

the    second,    third,    and    fourth    internodes    may 

begin  to  lengthen  long  before  the  first  has  attained 

its  full  growth.     Such  very  strong  buds  are  usually 

terminal ;  but  sometimes,  as  in  Lilac  (Fig.  86),  they 

are  the  uppermost  axillary,  which  take  the  place  of 

a  suppressed  or  abortive  terminal  bud. 

82.  Such  woody  stems,  developed  from  a  strong 
bud,  and  terminated  at  the  close  of  the  season's 
growth  by  a  similar  bud,  may  be  continued  from 
year  to  year  in  an  unbroken  series.   A  set  of  narrow 
rings  on  the   bark    (Fig.  85  a)    commonly  marks 
the  limit  of  each  year's  growth.     These  are  the 
scars  left  by  the  fall  of  the  scales  of  the  bud  ;  and 
these,  in  the  Horsechestnut,  and  in  other  trees  with 
large  scaly  buds,  may  be  traced  back  on  the  stem 
for  a  series  of  years,  growing   fainter  with   age, 
until  thej-  are  at  length  obliterated  by  the  action  of 

the  weather  and  the  distention  caused  by  the  increase  91 

of  the  stem  in  diameter.  The  same  is  the  case  with  the  more 
conspicuous  Leaf -scars,  or  marks  on  the  bark  left  by  the  separation 
of  the  leaf-stalk,  which  are  for  a  long  time  conspicuous  on  the 
shoots  of  the  Horsechestnut  (Fig.  85  5),  the  Magnolia  (Fig.  81), 
and  Hickory,  Fig.  91. 

83.  Ramification.     BRANCHES   (14-16)    are  secondary   stems 
developed  from  a  primary  one,  or  tertiary  ones  from  these,  and 
so  on.     Ultimate  or  small  ramifications  of  latest  order  are  some- 
times called  BRANCHLETS.     The  terminal  bud  continues  the  stem 
or  axis  which  bears  it.     Lateral  buds  give  rise  to  branches.1 
As  the  normal  lateral  buds  are  axillary  (75),  so  are  normal 
branches.     The  symmetry  or  arrangement  of  branches,  being 
that  of  the  buds  from  which  they  are  developed,  is  fixed  by  and 
follows  that  of  the  leaves.     When  the  leaves  are  alternate,  the 

1  Dichotomy  or  forking,  the  division  of  an  apex  into  two,  although  of  com- 
mon occurrence  in  the  lower  cryptogamous  plants,  occurs  so  rarely  and 
exceptionally,  if  at  all,  in  phaenogamous  plants  that  it  may  here  be  left 
out  of  view. 

In  phaenogamous  plants  only  the  ramification  of  axes  should  take  the 
name  of  branches.  That  is,  roots  and  stems  branch ;  and  the  term  may 
without  confusion  be  extended  to  hairs  and  all  TRICHOMES  (383)  when  com- 
pound, but  not  to  leaves  and  their  modifications. 

FIG.  91.  End  of  a  Hickory  branch  (Carya  alba),  with  a  strong  terminal  and  i 
axillary  buds*  "  • 


48  MORPHOLOGY  OF   STEMS. 

branches  will  be  alternate;  when  the  leaves  are  opposite,  and 
the  buds  develop  regularly,  the  branches  will  be  opposite,  &c. 
This  holds  in  fact  sufficiently  to  determine  and  exemplify  the 
plan  of  ramification  ;  but,  if  entirely  carried  out,  there  would  be 
as  many  branches  as  leaves.  This  could  rarely  if  ever  be,  even 
in  primary  ramification. 

84.  Non-development  of  Bads.     Some  of  the  buds  are  latent  or 
merely  potential,  that  is,  do  not  make  their  appearance  :  of  those 
which  do  appear  only  a  part  actually  grow  into  branches  ;  and 
of  these  some  are  apt  to  perish  at  an  early  stage.     In  our  trees, 
most  of  the  lateral  buds  generally  remain  dormant  for  the  first 
season :  they  appear  in  the  axils  of  the  leaves  early  in  summer, 
but  do  not  grow  into  branches  until  the  following  spring ;  and 
even  then  only  a  part  of  them  grow.     Sometimes  the  failure 
occurs  without  appreciable  order  ;  but  it  often  is  nearly  uniform 
in  each  species.     Thus,  when  the  leaves  are  opposite,  there  are 
usually  three  buds  at  the  apex  of  a  branch  ;  namely,  the  terminal, 
and  one  in  the  axil  of  each  leaf ;  but  it  seldom  happens  that  all 
three  develop  at  the  same  time.     Sometimes  the  terminal  bud 
continues  the  branch,  the  two  lateral  generally  remaining  latent, 
as  in  the  Horsechestnut  (Fig.  85)  ;  sometimes  the  terminal  one 
fails,  and  the  lateral  ones  grow,  when  the  stem  annually  becomes 
two-forked,  as   in   the  Lilac,  Fig.  86.     The  undeveloped  buds 
do  not  necessarily  perish,  but  are  ready  to  be  called  into  action 
in  case  the  others  are  checked.     When  the  stronger  buds  are 
destroyed,  some  that  would  else  remain  dormant  develop  in  their 
stead,  incited  by  the  abundance  of  nourishment,  which  the  for- 
mer would  have  monopolized.     In  this  manner  our  trees  are  soon 
reclothed  with  verdure,  after  their  tender  foliage  and  branches 
have  been  killed  by  a  late  vernal  frost,  or  consumed  by  insects. 
And  buds  which  have  remained  latent  for  several  years  occasion- 
ally shoot  forth  into  branches  from  the   sides   of  old   stems, 
especially  in  certain  trees. 

85.  Most  branches  springing  from  old  trunks,  however,  as  in 
Willows  and  Poplars,  especially  when  wounded  or  pollarded, 
originate   from  adventitious   buds    (77),   which   occur  without 
order.    So  also  when  accessory  buds  (76)  develop  into  branches, 
normal  symmetry  is  more  or  less  disturbed,  as  by  contiguous 
shoots  standing  directly  over  each  other  in  Tartarean  Honey- 
suckle, or  by  a  branch  far  out  of  the  axil  in  Walnuts  (Fig.  89) 
and  Honey-Locust,  Fig.  96. 

86.  Excurrent  and  Deliquescent  Stems.     Sometimes  the  primary 
axis  is  prolonged  without  interruption,  even  through  the  whole 
life  of  a  tree  (unless  accidentally  destroyed) ,  by  the  continued 


CHARACTERISTICS   AND    GROWTH.  49 

evolution  of  a  terminal  bud,  or  by  some  upper  strong  bud  which 
equally  becomes  a  leader,  —  forming  an  undivided  main  trunk, 
from  which  lateral  branches  proceed ;  as  in  most  Fir-trees. 
Such  a  trunk  is  said  to  be  excurrent.  In  other  cases,  the  main 
stem  is  arrested,  sooner  or  later,  either  by  flowering,  by  the 
failure  of  the  terminal  shoot,  or  by  the  more  vigorous  develop- 
ment of  some  of  the  lateral  buds  ;  and  thus  the  trunk  is  dissolved 
into  branches,  or  is  deliquescent,  as  in  the  White  Elm  and  most  of 
our  deciduous-leaved  trees.  The  first  naturally  gives  rise  to  coni- 
cal or  spire-shaped  trees  ;  the  second,  to  rounded  or  spreading 
forms.  As  stems  extend  upward  and  evolve  new  branches,  those 
near  the  base,  being  overshadowed,  are  apt  to  perish,  and  thus 
the  trunk  becomes  naked  below.  This  strikingl}'  occurs  in  the 
excurrent  trunks  of  Firs  and  Pines,  grown  in  forest,  which  seem 
to  have  been  branchless  to  a  great  height.  But  the  knots  in 
the  centre  of  the  wood  are  the  bases  of  branches,  which  have 
long  since  perished,  and  have  been  covered  with  a  great  number 
of  annual  layers  of  wood,  forming  the  clear  stuff  of  the  trunk. 

87.  Definite   and  Indefinite  Annual  Growth  of  Branches.     In 
man}*-  of  our  trees  and  shrubs,  especially  those  with  scaly  buds, 
the  whole  year's  growth  (except  on  certain  vigorous  shoots)  is 
either  already  laid  down  rudimentally  in  the  bud,  or  else  is  early 
formed,  and  the  development  is  completed  long  before  the  end 
of  summer ;  when  the  shoot  is  crowned  with  a  vigorous  terminal 
bud,  as  in  the  Horsechestnut  (Fig.  85)  and  Magnolia  (Fig.  81), 
or  with  the  uppermost  axillary  buds,  as  in  the  Lilac  (Fig.  86) 
and  Elm.     Such  definite  shoots  do  not  die  down  at  all  the  follow- 
ing winter,  but  grow  on  directly,  the  next  spring,  from  these 
terminal  or  upper  buds,  which  are  generally  more  vigorous  than 
those  lower  down.     In  other  cases,  on  the  contrary,  the  branches 
grow  onward  indefinitely,  until  arrested  by  the  cold  of  autumn  : 
the  buds  at  or  near  their  summit  are  consequently  young  and 
unmatured,  or  at  least  the  lower  and  older  axillary  buds  are 
more  vigorous,  and  alone  develop  into  branches  the  next  spring ; 
the  later-formed  upper  portion  most  commonly  perishing  from 
the  apex  downward  for  a  certain  length  in  the  winter.     The 
Rose  and  Raspberry,  and  among  trees  the  Sumac  and  Honey- 
Locust,  are  good  illustrations  of  this  sort ;    and  so  are  most 
perennial  herbs,  their  stems  dying  down  to  or  beneath  the  sur- 
face of  the  ground,  where  the  persistent  base  is  charged  with 
vigorous  buds,  well  protected  by  the  ground,  for  the  next  year's 
vegetation. 

88.  Many  of  the  details  and  applications  of  ramification,  of 
most  importance  in  morphology  and  descriptive  botany,  relate 


50  MORPHOLOGY  OP   STEMS. 

to  anthotaxy  or  inflorescence  (Chap.  V.),  which  has  its  own 
terminology.  But  some  of  its  terms  may  be  conveniently 
employed  in  the  description  of  ramification  unconnected  with 
flowering. 

§  2.   FORMS  OF   STEMS  AND  BRANCHES. 

89.  On  the  size  and  duration  of  the  stem  the  oldest  and  most 
obvious    division   of   plants  is  founded,   namely,    into   Herbs, 
Shrubs,  and  Trees. 

90.  Herbs  are  plants  in  which  the  stem  does  not  become 
woody  and  persistent,  but  dies  annually  or  after  flowering,  down 
to  the  ground  at  least.     The  difference  between  annual,  biennial, 
and  perennial  herbs  has  already  been  pointed  out  in  the  chapter 
on  the  root  (50-57) ,  and  the  gradations  between  them  indicated. 
Herbs  pass  into  shrubs  and  shrubs  into  trees  through  every  gra- 
dation.    The  following  definitions  are  therefore  only  general :  — 

91.  Undershrubs,  or  Sujffruticose  plants,  are  wood}*  plants  of 
humble  stature,  their  stems  rising  little  above  the  surface.     If 
less  decidedly  woody,  they  are  termed  Sujfrutescent. 

92.  Shrubs  are  wood}'  plants,  with  stems  branched  from  or 
near  the  ground,  and  less  than  five  times  the  height  of  a  man. 
A  shrub  which  approaches  a  tree  in  size,  or  imitates  it  in  aspect, 
is  said  to  be  Arborescent. 

93.  Trees  are  woody  plants  with  single  trunks,  which  attain 
at  least  four  or  five  times  the  human  stature.     Yet  the  name  of 
tree  is  not  to  be  denied  to  a  woody  plant  having  a  single  and 
stout  trunk  of  less  altitude ;  and  those  which  grow  in  a  bushy 
manner,  sending  up  a  cluster  of  stems  from  the  ground  to  the 
height  of  thirty  feet  or  more,  may  still  be  called  shrubs. 

94.  The  erect  position,  elevation  above  the  soil,  and  self-sup- 
port, are  normal  conditions  of  the  stem,  but  are  far  from  universal. 
And  certain  kinds  of  stem  or  branches  are  sufficiently  peculiar 
to  have  received  substantive  names  :  other  equally  peculiar  forms 
have  no  special  names.     There  are,  moreover,  certain  organs 
(such  as  spines  and  tendrils)  which  are  commonly  homologous 
(12)  with  stems,  but  not  always.     Two  kinds  of  erect  stems 
have  special  names  in  descriptive  botany. 

95.  Culm  is  a  name  applied  to  the  peculiar  closed-jointed  stem 
of  Grasses  and  Sedges,  whether  herbaceous,  as  in  most  Grasses, 
or  woody  or  arborescent,  as  in  the  Bamboo. 

96.  Candex  is  the  name  technically  applied  to  the  trunk  of 
Palms  (Fig.  126),  Tree-Ferns,  and   the  like,   consisting  of  a 
commonly  simple  column,  the  surface  beset  with  scales,  —  the 


SPECIAL  FORMS.  51 

bases  of  former  leaf-stalks,  —  or  marked  by  scars,  left  by  their 
fall.  This  name  was  used  by  botanists  anterior  to  Linnaeus  for 
any  tree-trunk,  but  is  now  used  for  the  peculiar  stems  above- 
mentioned  ;  also  for  the  persistent  base  of  a  stem,  otherwise 
annual,  which  throws  up  fresh  herbaceous  stems  or  stalks  from 
year  to  year.  Such  short  and  enduring  stems,  being  usually 
near  the  ground  or  under  it,  were  commonly  mistaken  for  roots. 
The  old  English  name  of  Stock  is  sometimes  used  in  botanical 
description  for  all  short  and  enduring  stems  of  this  sort, 
whether  rising  somewhat  above  or  concealed  beneath  the  surface 
of  the  soil. 

97.  A  Scape  is  a  stem  or  branch  which  rises  from  beneath  or 
near  the  surface  of  the  ground  and  bears  flowers,  but  no  proper 
foliage.     It  therefore  belongs  to  inflorescence.  (265.)     Scapes 
usually  spring  from  some  one  of  the  subterranean  forms  of  stem. 

98.  Of  stems  which  do  not  stand  upright  in  the  air  there  are 
various  modifications  and  gradations. 

99.  Scandent    or    Climbing   Stems  are  those  which  rise  by 
attaching  themselves  to  some  extraneous   support.      This  is 
effected  in  various  ways ;  in  some  by  the  action  of  the  stem 
itself,  in  others  by  that  of  organs  which  it  bears.1 

100.  Voluble  or  Twining  Stems,  or  Twiners,  are  those  which 
ascend  by  coiling  round  a  support,  which  must  accordingly  be 
comparatively  slender,  or  at  least  not  too  large.     Some  ascend 
by  coiling  "with  the  sun"  (that  is,  from  right  to  left  of  the 
observer  viewing  the  coil  from  the  outside 2) ,  as  the  Hop  ;  more, 

1  See  Darwin,  The  Movements  and  Habits  of  Climbing  Plants,  London 
and  New  York,  1875.    Also  the  earlier  paper  on  the  subject  in  Journal  of 
the  Linnean  Society,  ix.  1865. 

Note  that  in  North  America  climbing  plants  in  general  are  in  popular 
language  called  Vines  (e.  g.  Hop- Vine,  Grape- Vine,  Squash- Vine,  &c.),  a 
name  which  properly  belongs  to  Vitis  only. 

2  Dextrorse  and  Sinistrorse,  i.  e.  to  the  right  or  to  the  left,  are  almost  indis- 
pensable terms,  but  there  is  an  ambiguity  and  discrepancy  hi  their  use. 
Darwin  (in  Climbing  Plants,  above  referred  to)  seeks  to  avoid  this  by  usually 
employing  the  terms  "  with  the  sun,"  and  "  against  the  sun,"  phrases  which 
would  be  unmanageable  in  terminology.     The  writer  (in  Amer.  Jour.  Sci. 
ser.  3,  xiii.  391 )  suggested  Eutropic  for  the  former,  Antitropic  for  the  latter, 
to  be  used  in  case  it  is  preferred  to  evade  rather  than  to  encounter  the 
ambiguity.     Probably  the  terms  dextrorse  and  sinistrorse,  or  right  and  left, 
will  continue  in  use,  as  most  natural  and  convenient.     Now,  in  the  first 
place,  it  should  be  understood  that  a  plant,  or  at  least  a  plant's  axis,  having 
no  front  and  back,  can  have  no  right  and  left  of  its  own.     These  relations 
of  direction  must  refer  to  the  right  and  left  of  an  observer.     All  depends, 
accordingly,  upon  the  position  which  the  viewing  observer  is  supposed  to 
occupy  when  he  predicates  the  direction  of  the  turns  of  a  helix  or  of  the  over- 
lapping of  the  parts  of  a  bud.    Linnaeus  supposed  the  observer  to  view  the 


52  MOEPHOLOGY   OF   STEMS. 

by  coiling  in  the  opposite  direction,  as  the  Bean  (Phaseolus) , 
the  woody  Aristolochia  Sipho,  the  Morning  Glory  (Fig.  91°)  and 

other  Convolvu- 
laceae.  The  Dod- 
der, a  leafless  par- 
asitic plant  of  the 

latter  family,  not  only  gains  support  by  coiling 
on  the  stems  of  other  plants,  but  by  attachment, 
through  the  development  of  sucker- like  discs, 
along  the  whole  contiguous  surface.  (Fig.  77.) 
The  various  actions  through  which  plants  climb, 
and  the  attendant  phenomena,  are  physiological, 
and  will  be  treated  in  the  second  part  of  this 
Text  Book.  The  most  complete  and  satisfac- 
tory discussion  of  the  subject,  of  a  readable  sort, 
is  that  of  Darwin's  volume,  referred  to  in  a 
preceding  note. 

101.  Leaf-Climbers  are  those  in  which  support  is  gained  by  the 
action,  not  of  the  stem  itself,  but  of  the  leaves  it  bears  ;  in  most 
by  the  coiling  or  clasping  of  petioles,  as  in  Clematis,  Maurandia, 
Tropaeolum,  and  Solanum  jasminoides  (Fig.  235)  ;  in  some  by 
the  incurvation  of  leaf-blades  or  portions  of  them,  as  in  Adlu- 
mia  ;  or  by  an  extension  of  the  midrib  into  a  hook  or  short  ten- 
dril, as  in  Gloriosa ;  or  by  the  transformation  of  some  of  the 
blades  of  a  compound  leaf  into  hooks  or  tendrils,  as  in  Cobsea 
and  the  Pea. 

102.  Tendril-Climbers  (Fig.  92-95)  are  those  in  which  the 
prehension  is  by  a  tendril,  a  slender  filiform  body,  either  simple 
or  branched,  specially  adapted  to  the  purpose,  and  capable  of 
coiling,  either  to  secure  a  hold,  or  to  draw  the  stem  up  to  the 

coil  or  circle  from  the  inside ;  Mohl,  Palm,  Braun,  and  the  DeCandolles 
adopt  this,  and  the  latter  insist  on  it.  Such  authority  should  be  decisive, 
if  common  usage  and  popular  sense  went  along  with  it.  But  some  of  the 
botanists  following  Linnaeus  adopted  the  reverse  view ;  and  to  the  present 
writer,  as  to  Bentham  and  Hooker,  Darwin,  Eichler,  and  in  part  G.  Henslow, 
it  was  so  natural  to  view  the  coil  from  the  outside  that  we  without  concert 
adopted  this  position  and  mode  of  expression.  A  right-hand  coil,  or  one 
turning  to  the  right,  with  us,  is  one  the  turns  of  which  pass  from  the  left  to 
right  of  a  bystander  who  confronts  the  coil.  It  is  in  this  sense  that  a  com- 
mon screw  is  called  a  right-handed  screw,  and  that  the  right  bank  of  a  river 
ia  that  to  the  right  of  the  person  who  follows  the  course  of  the  stream.  So 
natural  is  this,  that  even  on  a  map  or  plate,  which  has  face  and  back,  and 
therefore  a  right  and  left  of  its  own,  the  figures  occupying  its  right  or  left 
portions  are  understood  to  be  those  which  are  toward  the  right  or  the  left 
hand  of  the  observer  who  stands  before  it. 

FIG.  91«.    Dextrorsely  twining  stem  of  Morning  Glory,  Ipomoea  purpuroa. 


SPECIAL  FOKMS.  53 

support.  In  certain  tendrils  the  attachment  to  the  support  is  by 
a  sucker- like  disc  at  the  apex,  as  in  the  Virginia  Creeper  or 
Ampelopsis,  Fig.  94. 

103.  Root-Climbers  are  those  in  which  the  stems  produce  aerial 
rootlets   (59a),   which  fix   themselves  to  a  supporting   surface 
along  which  the  stem  creeps  or  ascends.     In  this  way  Trumpet 
Creeper  (Tecoma  radicans) ,  Ivy,  and  Poison  Ivy  (Rhus  Toxi- 
codendron)  climb  extensively. 

104.  Stems  or  branches  which  neither  climb  nor  stand  upright 
may  have  their  direction  or  habit  of  growth  expressed  by  certain 
adjective  terms  ;  such  as 

Ascending  or  Assurgent,  when  they  rise  obliquely  upward ; 

Reclining,  'when  from  an  ascending  or  erect  base  the  upper 
part  recurves  and  trails ; 

Decumbent,  when  trailing  along  the  ground,  but  with  apex 
assurgent ; 

Procumbent  or  Prostrate,  when  lying  at  length  upon  the  ground  ; 

Repent  or  Creeping,  when  growing  prostrate  on  the  ground 
and  rooting  as  they  grow.  Also  applied  to  similar  stems  grow- 
ing under,  as  well  as  upon  the  surface  of  the  soil,  as  in  Couch- 
Grass  and  Mint,  Fig.  99. 

105.  A  Sucker  (Surculus}  is  an  ascending  stem  rising  from  a 
subterranean  creeping  base.     The  Rose  and  Raspberry  multiply 
freely  by  suckers.     Such  plants  are  easiest  to  propagate  "  by 
division." 

106.  A  Stolon  is  a  prostrate  or  reclined  branch  which  strikes 
root  at  the  tip,  and  then  develops  an  ascending  growth,  which 
becomes  an  independent  plant. 

107.  An  Offset  is  a  short  stolon  or  a  short  sucker.     Houseleek 
(Fig.  91*)   offers   a    familiar 

example.  By  offsets,  some 
herbs,  otherwise  annuals,  are 
continued  from  year  to  year  in 
a  vegetative  progeny  (Lobelia 
cardinalis,  &c.),  and  peren-i 
nials  may  thus  establish  colo- 
nies around  a  parent  individual. 

108.  A  Runner  (Flagellum) 
is  a  filiform  or  very  slender 

stolon,  naked  and  tendril- like  except  at  tip,  where  it  roots, 
develops  a  bud,  and  so  a  new  plant.  The  Strawberry  furnishes 
the  most  familiar  example. 

FIG.  91*.    Houseleek  (Semperviytun  tectorum)  with  oflseta. 


54  MORPHOLOGY   OF   STEMS. 

109.  The  two  following  are  organs  which  may  be  of  axial 
nature,  or  may  not.  This  may  ordinarily  be  determined  by  posi- 
tion. Any  direct  continuation  of 
stem  or  branch  must  be  of  axial 
nature,  that  is,  of  the  nature  of  stem ; 
and  the  same  is  true  of  whatever 
primarily  develops  in  the  axil  of  a 
leaf.  Conversely,  whatever  subtends 
a  lateral  axis  or  branch  may  be  taken 
for  a  leaf  or  foliar  production,  being 
in  the  place  of  such. 

110.  A  Tendril,  a  thread-shaped 
and  leafless  body,  capable  of  coiling 
spirally,  and  used  for  climbing  (102) , 
is  homologous  with  stem  in  Grape- 
vines (Fig.  92)  ;  for  the  uppermost 
tendril  is  seen  to  be  a  direct  continu- 
ation of  the  stem.  The  small  bud 
which  appears  in  the  axil  of  the 
uppermost  leaf  will  in  its  growth 
produce  another  internode  and  leaf, 
or  some  species  more  than  one, 
but  will  terminate  in  a  similar 
tendril:  the  present  terminal  tendril  will  have  then  become 
lateral  and  opposite  the  leaf,  like  the  three  in  the  lower  part  of 


FIG.  92.  End  of  a  shoot  of  the  Grape-vine,  with  young  tendrils:  a  sympodial 
•tern.  (See  note.) 

FIG.  93.  A  portion  of  a  stem  of  Ampelopsis  quinquefolia,  or  Virginia  Creeper,  with 
a  leaf  and  a  tendril. 

FIG.  94.  Ends  of  the  latter,  enlarged,  showing  the  expanded  tips  or  discs  by  which 
they  cling. 


SPECIAL  FORMS.  55 

the  figure.1  The  tendrils  of  Virginia  Creeper  (Fig.  93)  are  of  the 
same  nature  and  position.  But,  instead  of  laying  hold  by  a  coiling 
of  the  tip,  when  it  has  reached  any  solid  surface,  such  as  a  wall 
or  tree-trunk,  the  tip  expands  into  an  adhesive  disc,  which  forms 
a  secure  attachment.  (Fig.  94.)  In  a  related  plant,  Vitis  (Cissus) 
tricuspidata  of  Japan,  these  disks  terminate  the  branches  of  ver}r 
short  tendrils  :  consequently  the  shoots  as  they  grow  are  at 
once  applied  closety  and  secured  finnty  to  the  surface  of  the  sup- 
port, —  an  admirable  adaptation  for  climbing  walls  and  trunks. 


111.  The  simple  tendril  of  a  Passi9n- 
flower,  being  in  the  axil  of  a  leaf  (that 
is  in  the  position  of  a  branch) ,  is  also  of 
axial  nature :  it  is  a  leafless  and  simple 
branch,  composed  of  one  long  and  slen- 
der internode,  devoted  to  the  purpose  of 
climbing.  Fig.  95  shows  in  all  stages  the 
admirably  active  tendrils  of  Passiflora 
sicyoides.  This  is  a  Mexican  species, 
remarkable  for  the  rapidity  and  freedom 
with  which  the  tendrils  move.  The  lowest 
tendril  in  the  figure  is  attached  and 
coiled :  the  next  is  free  and  coiled  in 
one  helix :  the  third  is  outstretched 
and  seeking  a  support.  For  tendrils 
which  are  not  homologous  with  stems, 
see  Sect.  IV.  228. 
112.  A  Spine  or  Thorn  (Fig.  96,  97)  is  usually  a  branch  or 
the  termination  of  a  stem  or  branch,  indurated,  leafless,  and 
attenuated  to  a  point.  The  nature  of  spines  is  manifest  in  the 
Hawthorn  (Fig.  97),  not  only  by  their  position  in  the  axil  of  a 
leaf,  but  often  by  producing  imperfect  leaves  and  buds.  And 
in  the  Sloe,  Pear,  &c.,  many  of  the  stinted  branches  become 
spinose  or  spinescent  at  the  apex,  tapering  off  gradually  into  a 
rigid  and  leafless  point,  thus  exhibiting  every  gradation  between  a 
spine  and  an  ordinary  branch.  These  spinose  branches  are  less 

1  This  forms  what  is  called  a  Sympodium  or  Sympodial  stem,  which  is  mor- 
phologically made  up  of  a  series  of  superposed  branches.  (See  Chapter  V. 
281,282.)  In  contradistinction,  a  stem  formed  by  the  continued  development 
of  a  terminal  bud  is  Monopodial  or  a  Monopodium.  Fig.  95  is  an  example. 

FIG.  95.  Leafy  shoot  of  Passiflora  sicyoides,  of  Mexico,  with  fixed  and  coiled,  free 
and  full  grown,  and  forming  tendrils. 


56 


MORPHOLOGY    OF   STEMS. 


liable  to  appear  on  the  cultivated  tree,  when  duty  cared  for,  such 
branches  being  thrown  mostly  into  more  vigorous  growth.     In 

the  Hawthorn,  the  spines 
spring  from  the  main  axillary 
bud,  while  accessory  buds 
(76),  one  on  each  side,  ap- 
pear, and  grow  the  next  sea- 
son into  ordinary  branches. 
In  the  Honey-Locust,  it  is 
the  uppermost  of  several  ac- 
cessory buds,  placed  far  above 
the  axil,  that  develops  into 
the  thorn  (Fig.  96) .  Here  the 
spine  itself  usually  branches, 
and  sometimes  becomes  ex- 
tremely compound. 

113.  For  spines  which  are 
homologous  with  leaves,  or 
parts  of  a  leaf,  see  Sect.  IV. 
227°.  Prickles,  such  as  those 
of  Brambles  and  Roses,  are 
superficial  outgrowths  from 
the  bark,  of  a  different  nature 
(383),  and  of  small  morpho- 
logical signification. 

114.  Subterranean  Stems  are  hardly  less    diverse    than    the 
aerial.     They  are  classed  as  RHIZOMES,   TUBERS,  CORMS,  and 
BULBS,  the  forms  passing  one  into  another  by  gradations. 

115.  Rhizoma  (Rhizome,  or  in  English  ROOTSTOCK)  is  a  gen- 
eral name  for  any  horizonal  or  oblique  perennial  stem,  which  lies 
on  the  ground  or  is  buried  beneath  its  surface.     It  sends  off 
roots  of  a  fibrous  or  slender  sort  wherever  it  rests  on  or  is  cov- 
ered by  the  soil,  and  usually  produces  from  its  apex  some  kind 
of  aerial  stem,   either  leafy  or  as  a  flower-stalk   (scape,  97), 
which  rises  into  the  air  and  light.      Before  morpholog}T  was 
understood,  rootstocks  were  called  creeping  roots,  scaly  roofs,  &c. 
Some  are  slender,  such  as  those  of  Mints   (Fig.  99),  of  most 
Sedges  (Fig.  98),  and  of  Couch-Grass.     Their  cauline  nature  is 
evident  from  their  structure  and  appearance ;  their  nodes  and 
internodes  are  well  marked,  the  former  bearing  leaves  reduced  to 

FIG.  96.  Branching  thorn  of  the  Honey-Locust  (Gleditschia),  an  indurated  branch 
developed  from  an  accessory  bud  produced  above  the  axil.  a.  Three  buds  under  the 
base  of  the  leaf-stalk,  brought  to  view  in  a  section  of  the  stem  and  leaf-stalk  below. 

Fid  97.  Thorn  of  the  Cockspur  Thorn,  developed  from  the  central  of  three  axillary 
buds;  one  of  the  lateral  buds  is  seen  at  its  base. 


SPECIAL   FORMS. 


57 


scales  y  and  the  advancing  apex  rises  at  length  into  an  ordinary 
stem,  while  the  opposite  and  older  end  gradually  dies  away.  A 
bud  forms  in  the  axil  of  each  scale-like  leaf,  or 
in  some  of  them  ;  roots  proceed  from  the  nodes 
in  preference  ;  the  destruction  of  the  ascending 
stem  only  brings  these  buds  into  activity ;  and 
the  cutting  or  tearing  of  the  rootstock  into 
pieces  by  the  hoe  or  plough  merely  hastens  the 
establishment  of  as  many  new  plants,  each  with 
roots,  bud,  and  a  small  store  of  nourishment 
ready  provided.  It  is  this  which  makes  Couch- 
Grass  or  Quick-Grass  (Triticum  repens)  very 


troublesome  to  the  agriculturist ;  and  the  Nut-Grass    (Cyperus 
rotundus,  var.  Hydra)  of  the  Southern  Atlantic  States  is  even 


more  so,  portions  of  its  rootstock  being  tuberiferous,  i.  e.  en- 
larged into  a  tuber  which  contains  a  supply  of 
concentrated  nourishment  to  feed  the  growth. 

116.  Thickened  rootstocks  are  common; 
nourishing  matter,  elaborated  in  the  leaves 
above,  being  accumulated  in  them,  just  as 
it  is  in  thickened  roots,  and  for  the  same  pur- 
pose. (53-55.)  Such  are  the  so-called  roots  of  Sweet-Flag,  of 
Ginger,  of  Iris  or  Flower-de-Luce  (Fig.  216),  of  Bloodroot,  of 
Solomon's  Seal  (Fig.  100),  &c.  These  grow  after  the  manner 
of  ordinary  stems,  advancing  from  year  to  year  by  the  annual 
development  of  a  bud  at  the  apex,  and  emitting  roots  from  the 
under  side  or  the  whole  surface.  Thus  established,  the  older 


FIG.  98.  Slender  rhizoma  of  Carex  arenaria,  of  Europe,  which  binds  shifting  sands 
of  the  sea-shore. 

FIG.  99.    Rootstocks,  or  creeping  subterranean  branches,  of  the  Peppermint. 

FIG.  99°.  A  piece  of  the  rootstock  of  the  Peppermint,  enlarged,  with  its  node  or  joint, 
and  two  axillary  buds  ready  to  grow. 


58 


MORPHOLOGY   OF   STEMS. 


portions  die  and  decaj-  as  corresponding  additions  are  made  to 
the  opposite  growing  extremity.     Each  year's  growth  is  often 


marked  conspicuously,  sometimes  by  a  strong  contraction  where 
the  interruption  took  place,  as  in  certain  species  of  Iris  (Fig.  216) ; 

or  by  the  circular  im- 
pressed scar  (likened  to 
the  impression  of  a  seal) 
in  Solomon's  Seal ;  this 
being  the  place  where 
the  annual  aerial  stem, 
bearing  the  vegetation, 
separated  in  autumn 
from  the  perennial  rhi- 
zoma.  The  numerous 
slender  lines  encircling  the  rootstock  are  the  scars  left  after  the 
decay  of  the  scale-like  leaves  or  bud-scales,  such  as  are  seen  at 
the  young  and  growing  end  of  the  rootstock. 
The  rootstock  of  Diphylleia,  of  the  Alleghany 
Mountains  (Fig.  101),  is  similar;  but  the 
yearly  growths  are  so  exceedingly  short  that 
they  become  vertical,  the  bud  of  each  year 
is  close  to  the  stalk  of  the  year  preceding, 
and  the  scars  marking  previous  growths  are 
in  contact.1  Trillium  makes  a  short  and 
mostly  vertical  rootstock,  which,  when  it 
remains  simple  and  dies  away  promptly 
below  (as  in  Fig.  102),  comes  nearly  within 
102  VVI  '  the  definition  of  a  conn.  But  in  several 


1  The  rootstock  in  Polygonatum  and  Diphylleia  is  a  sympodium  (110,  note), 
the  terminal  bud  developing  yearly  the  growth  above  ground  and  perishing 

FIG.  100.  Rootstock  of  Polygonatum  or  Solomon's  Seal,  with  the  terminal  bud,  the 
base  of  the  stalk  of  the  season,  and  three  scars  from  which  the  latter  has  separated  in 
ai  many  former  years. 

FIG.  101.  Rhizoma  of  Diphylleia  cymosa,  showing  six  years'  growth,  and  a  bud  for 
the  seventh :  a,  the  bud :  b,  base  of  the  stalk  of  the  current  year :  c,  scar  left  by  the 
decay  of  the  annual  stalk  of  the  year  before ;  and  beyond  are  the  scars  of  previous  years. 

FIG.  102.    Shoot  and  young  rootstock  of  Trillium  erectum,  with  only  terminal  bud. 


SPECIAL  FORMS. 


59 


species,  and  in  older  individuals,  it  is  longer,  often  oblique,  and 

branching,  and  bears  the  scars  from  which  the  annual  aerial 

growths  have  separated.1    Nymphaea  odorata, 

the  sweet-scented  white  Water  Lily,  grows  by 

very  long,  stout,  and  simple  rootstocks.     In 

N.  tuberosa  the  sides  of  the  rootstock  produce 

short  lateral  branches  or  tubers. 

117.  A  Tuber  may  be  morphologically  char- 
'acterized  as  a  short  thickened  rhizoma  on  a 
slender  base,  or  a  rootstock  some  portion  of 
which — mostly  a  terminal  portion  and  involv- 
ing several  nodes  —  is  thickened  by  the  depo- 
sition of  nourishing  matter.  A  potato  and  a 
Jerusalem  artichoke  are  typical  examples 
(Fig.  104-107)  ;  and  the  diiference  between 
these  subterranean  branches  and  the  roots  which  they  may  bear 
is  very  obvious.  Their  eyes  are  axillary  buds  ;  the  leaves  which 
subtend  them  are  plainly  dis-  m 
cernible,  in  the  form  of  short  and 
closely  appressed  scales.  In  the 
attempt,  occasionally  seen,  to 
form  axillary  tubers  above- 
ground  by  the  Potato-plant,  the 
leafy  nature  of  the  scales  is 
evidenced.  (Fig. 
105.)  By  heaping 
the  soil  around 
the  stems,  the- 
number  of  tuber-  \ 
iferous  branches 
may  be  in- 
creased. The  number  of  nodes  and  internodes  involved  in  a 
tuber  may  be  many  or  few.  There  is  one  instance  of  what  may 

in  autumn,  to  be  renewed  by  an  axillary  bud,  which  makes  its  subterranean 
growth  and  the  rudiments  of  the  aerial  in  early  summer. 

1  This  rhizoma  is  a  monopodium,  being  continued  year  by  year  by  the 
terminal  bud,  and  the  aerial  stem  or  stems  sent  up  in  spring,  bearing  the 
whorl  of  leaves  and  blossom,  are  axillary  branches. 

FIG.  103.  An  older  and  longer  one  of  the  same  species,  showing  branches,  scars  left 
by  former  leaf-  and  flower-bearing  stems :  also  at  tip  (stripped  of  the  covering  scales),  the 
bases  of  two  such  stems  of  the  season,  and  the  terminal  bud  between  them,  for  the  con- 
tinuation of  the  growth  of  the  rootstock,  &c.,  the  next  season. 

FIG.  104.  Base  of  stem  of  Helianthus  tuberosus,  or  Jerusalem  Artichoke,  developed 
from  a  tuber,  and  producing  a  second  generation  of  tubers. 

FIG.  105.  Monstrosity  of  a  Potato-plant,  with  an  axillary  bud  developing  into  some- 
thing between  a  bulblet  and  a  tuber ;  the  scales  represented  by  obvious  leaves.  (From 
the  Gardeners'  Chronicle.) 


60 


MORPHOLOGY   OF   STEMS. 


be  called  a  Monomerous  tuber,  namely  in  Nelumbium  luteum 
(Fig.  108"),  where  it  consists  of  a  single  thickened  internode  of 


an  aquatic  runner,  which  is  accordingly  quite  destitute  of  scales 
or  buds.     The  growth   proceeding   from  this   simple   tuber  is 

necessarily  from  a  bud  of 
the  node  at  its  apex,  whence 
also  a  cluster  of  roots  is 
produced.  Of  a  somewhat 
similar  nature  are  the  con- 
catenate tubers  of  Apios 
tuberosa  (several  of  which 
IDS"  are  strung  as  it  were  upon 

a  long  filiform   axis),  the  tubers  not  unfrequently  being  mo- 
nomerous,  although  the  larger  ones  are  not  so. 

117°.  Tubercles,  as  they  may  be  termed,  are  of  a  mixed  or 
ambiguous  character  between  tubers  and  tuberous  roots.  A  good 
example  of  the  latter  is  afforded  by  Dahlia-roots.  (Fig.  68.) 
They  yield  their  nourishing  substance  to  growing  buds  on  the 
stem  above,  but  do  not  themselves  normally  produce  even 

FIG.  106.  Forming  potatoes  in  various  stages.  107.  One  of  the  younger  ones  en- 
larged. 108.  Section  of  a  small  portion  passing  through  an  eye,  or  bud,  more  enlarged. 

FIG.  108o.  A  monomerous  (i.  e.  one-membered)  tuber  of  Nelumbium  luteum,  formed 
of  a  single  internode. 


SPECIAL  FORMS.  61 

adventitious  buds.  Sweet  potatoes  (55),  although  equally 
roots,  do  produce  adventitious  buds,  especially  from  near  the 
upper  end.  The  somewhat  similar  tubercles  or  tumefied  roots 
of  certain  Orchises  and  other  plants  of  the  same  tribe,1  definite 
in  number  and  shape,  and  sometimes  imitating  a  corm,  are 
charged  with  a  bud  at  the  upper  end,  near  their  origin.  Ap- 
parently, the  origin  is  a  bud  from  the  base  of  the  parent 
stem,  which  bud  directly  forms  a  tumefied  short  root  from  its' 
very  base.2 

118.  A.  Corm  (Cormus)  is  to  be  compared  on  the  one  hand 
with  a  short  rootstock  or  tuber,  on  the  other  with  a  bulb.     It 
is  a  subterranean   fleshy  stem,  of  rounded  or  depressed  figure 
and  solid  texture.     Some  of  its  buds  grow  into  new  corms,  and 
these,  upon  the  death  of  or  separation  from  the  parent,  become 
new  individuals  :  some  develop  above  ground 

the  vegetation  and  the  blossoms  of  the  season. 
A  good  type  of  corm  is  that  of  Cyclamen 
(Fig.  109),  in  which  the  very  base  of  the 
seedling  stem  grows  fleshy,  and  widens  from 
year  to  year,  but  hardly  at  all  lengthens,  and 
so  becomes  far  broader  than  high,  or  de- 
pressed. As  the  main  bulk  belongs  to  the 
first  internode,  or  caulicle,  the  buds  from  which  the  yearly 
growths  of  leaves  and  flower-stalks  spring  are  at  the  centre  of 
the  summit  or  upper  surface,  the  roots 
from  the  lower,  and  the  sides  seldom  pro- 
duce any  buds.  The  corm  of  Indian 
Turnip  (Arisaema  triphyllum,  Fig.  110) 
is  somewhat  similar,  but  it  sends  up  a 
single  stout  stem,  and  the  roots  spring 
from  around  the  base  of  this.  These  are 
completely  naked  corms. 

119.  But  in  Crocus  (Fig.  Ill,  112),  Colchicum  (Fig.  117),j 
Gladiolus,  and  the  like,  the  sheathing  bases  of  one  or  two  leaves' 
enclose  the  corm  with  a  membranous-scaly  coat,  giving  it  exactly 
the  appearance  externally  of  a  coated  bulb.    Such  have  been  not 
inappropriately  named  solid  bulbs.     In  common  parlance,  they 
will  doubtless  continue  to  be  called  bulbs,  and  even  in  popular 

1  Not,  however,  such  as  those  of  Aplectrum,  Tipularia,  etc.,  which  are 
genuine  corms  or  tubers. 

2  Irmisch,  Beitr.  Biol.  &  Morphol.  Orchid.  1853,  fide  Duchartre,  £le'm. 
Bot.  278. 

FIG.  109.    Depressed  corm  of  Cyclamen. 

FIG.  110.    Corm  of  Indian  Turnip,  Arisssma  triphyllum. 


62 


MORPHOLOGY   OF   STEMS. 


botanical  descriptions.  In  fact,  while  they  differ  from  naked 
conns  in  having  some  investment,  they  differ  from  true  bulbs 
only  in  the  greater  size  of  the  solid  axis  and  the  fewness  of 

the  investing  scales ; 
the  stem  or  solid  body 
making  the  greater  part 
of  the  corm,  but  a  very 
small  part  of  a  proper 
bulb.  There  are,  more- 
over, all  gradations  be- 
112  tween  the  two. 

120.  A  Bulb,  as  compared  with  a  corm,  may  be  said  to  be  an 
exceedingly  abbreviated  stem,  reduced  to  a  flat  plate,  from  the 
lower  face  of  which  roots  are  produced,  from  the  upper  face, 
leaves  in  the  form  of  scales ;  these  scales  being  either  reduced 
and  thickened  leaves  or  the  thickened  bases  of  ordinary  leaves. 
Compared  with  buds  (73),  it  is  a  very  fleshy  bud,  usually  large 
and  subterranean,  the  axis  of  which  never  elongates.  It  is  a 


114  115  116 

provision  for  future  growth,  the  stored  nourishment  of  which  Is 
deposited  in  the  leaves,  or  the  homologues  of  leaves,  instead  of 
in  the  stem. 

PIG.  111.  Conn  of  Crocus,  the  few  thin  enveloping  scales  removed,  showing  their 
scars,  which  mark  the  nodes,  the  shrivelled  vestige  of  the  last  year's  corm  at  the  base, 
and  buds  developing  into  new  ones  on  various  parts  of  its  surface.  112.  Vertical  section 
of  a  similar  corm,  with  a  terminal  and  one  lateral  bud. 

FIG.  113.    Section  of  a  tunicated  bulb  of  the  Onion. 

FIG.  114.  Vertical  section  of  the  bulb  of  the  Tulip,  showing  its  stem  or  terminal 
bud  (c)  and  two  axillary  buds  (6,  ft). 

FIG.  115.    Bulb  of  a  Garlic,  with  a  crop  of  young  bulbs. 

FIG.  116.    Vertical  section  of  the  conn  of  a  Crocus:  a.  new  buds. 

FIG.  117.  Vertical  section  of  the  conn  of  Colchicum  (6),  with  the  withered  conn  of 
the  preceding  (a),  and  the  forming  one  (c)  for  the  ensuing  year. 


SPECIAL  FOBMS. 


63 


121.  A  Tunicated  or  Coated  Bulb  (Fig.  113-115)  is  one  in 
which  the  scales  are  broad  and  completely  enwrapping,  forming 
concentric   coatings.     These  are  thickish  when  fresh,  but  thin 
when  exhausted  and  dry,  as  in  the  Onion,  Garlic,  and  Tulip. 

122.  A  Scaly  Bulb 
has    the   bulb-scales 
comparatively     nar- 
row, thick,  and  small, 
imbricated,   but   not 
severally  enwrapping 
each  other.    That  of 
the  Lily  is  the  most 
familiar    and     char-, 
acteristic     example.- 

(Fig.  118,  119.)  "8 

123.  Bulblets  are  small  aerial  bulbs,  or  buds  with  fleshy  scales, 
which  arise  in  the  axils  of  the  leaves  of  several  plants,  such  as 
the   common  Lilium  bulbiferum  and 

L.  tigrinum,  the  Tiger  Lilies  of  the 
gardens  (Fig.  120).  Here  they  ap- 
pear during  the  summer  as  axillary 
buds:  they  are  at  length  detached, 
and  falling  to  the  ground  strike  root, 
and  grow  as  independent  plants.  In 
the  common  Onion,  and  in  many  other 
species  of  Allium,  similar  bulblets 
take  the  place  of  flower-buds  in  the 
umbel.  Bulblets  plainly  show  the  identity  of  bulbs  with  buds. 

124.  All  these  extraordinary,  no  less  than  the  ordinary,  forms 
of  the  stem,  grow  and  branch,  or  multiply,  by  the  development 
of  terminal  and  axillary  buds.     This  is  perfectly  evident  in  the 
rhizoma  and  tuber,  and  is  equally  the  case  in  the  corm  and  bulb. 
The  stem  of  the  bulb  is  usually  reduced  to  a  mere  plate  (Fig. 
114 a),  which  produces  roots  from  its  lower  surface,  and  leaves 
or  scales  from  the  upper.     Besides  the  terminal  bud  (c) ,  which 
usually  forms  the  flower-stem,  lateral  buds  (5,  b)  are  produced 
in  the  axils  of  the  leaves  or  scales.     One  or  more  of  these  may 
develop  as  flowering  stems  the  next  season,  and  thus  the  same 
bulb  survive  and  blossom  from  year  to  year ;  or  these  axillary 
buds  may  themselves  become  bulbs,  feeding  on  the  parent  bulb, 
which  in  this  way  is  often  consumed  by  its  own  offspring,  as  in 

FIG.  118.   Scaly  bulb  of  Canada  Lily,  Liliuin  Canadense,  after  flowering.    119.  Ver- 
tical section  of  same,  showing  two  new  young  bulbs  within. 

FIG.  120.    Bulblets  in  the  axil  of  the  cauliue  leaves  of  Tiger  Lily. 


64  MORPHOLOGY   OF   STEMS. 

the  Garlic  (Fig.  115)  ;  or,  finally  separating  from  the  living 
parent,  just  as  the  bulblets  of  the  Tiger  Lily  fall  from  the  stem, 
they  may  form  so  many  independent  individuals.  So  the  corm 
of  the  Crocus  (Fig.  Ill,  112)  produces  one  or  more  new  ones, 
which  feed  upon  and  exhaust  it,  and  take  its  place ;  and  the 
next  season  the  shrivelled  remains  of  the  old  corm  ma}'  be  found 
underneath  the  new.  The  corm  of  Colchicum  (Fig.  117)  pro- 
duces a  new  bud  on  one  side  at  the  base,  and  is  consumed  by  it 
in  the  course  of  the  season  ;  the  new  one,  after  flowering  by  its 
terminal  bud,  is  in  turn  consumed  by  its  own  offspring ;  and 
so  on.  The  figure  represents  at  one  view,  a,  the  dead  and 
shrivelled  corm-  of  the  year  preceding ;  5,  that  of  the  present 
season  (in  a  vertical  section)  ;  and,  c,  the  nascent  bud  for  the 
growth  of  the  ensuing  year. 

125.  Condensed  Steins,  homologous  with  conns,  tubers,  &c., 
and  similar  in  mode  of  growth,  but  above  ground,  and  multiply- 
ing in  the  same  ways,  are  not  uncommon.  The  Cactus  family  is 
mainly  composed  of  such  forms,  of  flat-  or  round-jointed  Prickly 
Pears  (Opuntia) ,  fluted  or  angled  columns  (Cereus) ,  and  glob- 
ular Melon-Cactus,  Mamillaria,  and  Echinocactus.  The  latter 
types,  which  completel}'  imitate  corms,  are  the  most  consolidated 
forms  of  vegetation.  While  ordinary  plants  are  constructed  on 
the  plan  of  great  expansion  of  surface,  these  present  the  least 
possible  amount  of  surface  in  proportion  to  their  bulk,  their 
permanent  spherical  figure  being  that  which  exposes  the  smallest 
portion  of  their  substance  to  the  air.  Such  plants  are  evidently 
adapted  to  very  dry  regions  ;  and  in  such  only  are  they  naturally 
found.  Similarly,  bulbous  and  corm-bearing  plants,  and  the 
like,  are  a  form  of  vegetation  which  in  the  growing  season  may 
in  the  foliage  expand  a  large  surface  to  the  air  and  light,  while 
during  the  period  of  rest  the  living  vegetable  is  reduced  to  a 
globular  or  other  form  of  the  least  surface  ;  and  this  is  protected 
by  its  outer  coats  of  dead  and  dry  scales,  as  well  as  by  its  subter- 
ranean situation ;  —  thus  exhibiting  another  and  very  similar 
adaptation  to  a  season  of  drought.  And  such  plants  mainly 
belong  to  countries  (such  as  Southern  Africa,  and  the  interior 
of  Oregon  and  California)  which  have  a  long  hot  season,  during 
which  little  or  no  rain  falls,  when,  their  stalks  and  foliage  above 
and  their  roots  beneath  being  early  cut  off  by  drought,  the  plants 
rest  securely  in  the  corm-like  forms  to  which  they  are  reduced, 
and  retain  their  moisture  with  great  tenacity  until  the  rainy  season 
returns.  Then  they  shoot  forth  leaves  and  flowers  with  wonderful 
rapidity,  and  what  was  perhaps  a  desert  of  arid  sand  becomes 
green  with  foliage  and  gay  with  blossoms,  almost  in  a  day. 


SPECIAL  POEMS.  65 

126.  Stems  scrying  the  purpose  of  foliage,  Phyllocladia.    Most 
of  these  condensed  and  permanent  stems  are  illustrations  of 
this,  their  green  rind  doing  duty  for  leaves,  which 

are  either  absent,  or  transient,  or  reduced  to 
spines  or  other  organs  not  effective  as  foliage. 
In  the  flat  and  broad-jointed  species  of  Opuntia, 
and  still  more  in  PhyUocactus  and  Epiphyllum,  the 
forms  assumed  give  a  considerable  surface  of  green 
rind,  which  well  answers  the  purpose  of  leaves. 
Flattened  stems  or  branches  of  the  same  sort  and 
economy  not  seldom  occur  in  other  than  fleshy  or 
succulent  plants  (such  as  the  Cactuses)  ;  some- 
times accompanied  by  a  certain  number  of  real 
foliage-leaves,  but  these  more  or  less  transient, 
as  in  Bossiaea  and  Carmichaelia  among  Legu- 
minous shrubs,  and  Muhlenbeckia  platyclada,  now 
in  common  cultivation  (Fig.  121)  ;  sometimes 
with  all  the  leaves  reduced  to  small  and  function- 
less  scales,  as  in  the  Xylophylla  («'.  e.  wooden- 
leaved)  section  of  Phyllanthus,  and  in  Phyllo- 
cladus  (New  Zealand  and  Tasmanian  trees  of 
the  Yew  family) .  In  all  these,  the  cauline  nature 
is  manifest  by  the  continuous  or  proliferous 
growth,  by  the  marked  nodes  and  internodes, 
and  often  by  the  bearing  of  flowers. 

127.  Cladophylla  (literally,  branch-leaves)  are  more  ambigu- 
ous in  character.     The  most  familiar  examples  are  found  in  the 
peculiar  foliage  of  Ruscus,  Myrsiphyllum,  Asparagus,  and  in 
some  other  genera  of  the  same  family.     In  these  the  primary  or 
proper  leaves  of  the  shoots  are  little  scales,  one  to  each  node,  and 
quite  functionless.     From  the  axil  of  each  is  immediately  pro- 
duced a  body  answering  in  all  respects  to  the  blade  of  a  leaf, 
both  in  appearance  and  in  office.     They  also  accord  with  leaves 
in  being  expanded  horizontally,  although  they  take  a  twist  which 
brings  them  more  or  less  into  a  vertical  position,  in  the  manner 
of  phyllodia  (that  is,  of  leaf-stalks  assuming  the  form  and  office 
of  leaf- blades,  217) ;  wherefore  they  may  be  regarded  as  the 
first  and  only  leaf  of  an  axillary  branch  with  the   internode 
under  the  leaf  wholly  undeveloped  and  no  further  growth  ever 
taking  place.     But,  on  the  other  hand,  their  anatomical  structure 
is  said  to  be  that  of  stems  rather  than  of  leaves.     Moreover, 


FIG.  121.     Foliiform  branch  of  Muhlenbeckia  platyclada,  growing  from  the  apex, 
bearing  a  small  and  transient  leaf  at  some  nodesj  also  a  flower  or  two. 


66  MORPHOLOGY   OF   STEMS. 

the  cladophyll  of  Ruscus  (called  Butcher's  Broom  in  England, 
Fig.  123)  not  only  becomes  firm,  hard,  and  spiny-tipped,  but 
it  exhibits  the  character  of  a 
branch  by  bearing  flowers  on  the 
middle  of  one  face,  in  the  axil  of 
a  little  bract.  Under  this  view 
such  a  cladophyll  would  seem  to 
be  a  flattened  branch  of  two  in- 
ternodes,  or  else  of  one  internode 
with  a  flower-stalk  adnate  to  it. 
In  Myrsiphyllum  (a  South  Afri- 
can climber,  commonly  cultivated 
under  the  erroneous  name  of 
Smilax,  Fig.  122),  the  cladophyll 
is  wholly  leaf-like  in  appearance 
as  well  as  in  function,  and  it  never  bears  either  scale-leaf 
or  blossom  ;  but  the  flowers  are  on  slender  stalks  from  buds  out 
of  the  same  axil.  (See  Dickson  in  Trans.  Bot.  Soc.  Edinb. 
xvi.,  and  Van  Tieghem,  Bull.  Bot.  Soc.  France,  xxxi.,  for  a 
discussion  of  the  nature  of  cladophylla.) 

128.  To  all  such  leaves  or  imitations  of  leaves,  Bischoff  has 
given  the  name  PHYLLOCLADIA,  sing.  PHYLLOCLADIUM.  To  those 
definitely  restricted  to  one  internode,  and  which  so  closely 
counterfeit  leaves,  Kunth  gave  the  name  of  CLADODIA,  sing. 
CLADODIUM.  The  best  common  name  for  all  productions  which 
imitate  leaves  would  have  been  that  of  phyllodium 
(meaning  simply  a  leaf-like  bod}')  ;  but  that  term 
was  first  applied  and  is  restricted  to  the  case 
of  a  petiole  imitating  the  blade  of  a  leaf.  The 
name  Phyllocladium  (meaning  a  leaf-like  branch) 
may  properly  be  retained  for  the  whole  series  of 
leaf-like  bodies  here  described.  But  for  those  of. 
the  preceding  paragraph,  which  are  so  peculiarly 
leaf-like,  Kunth's  name  of  Gladodium  (i.  e.  a 
branch-like  body)  is  false  in  meaning,  and  may 
be  replaced  by  that  of  CLADOPHYLLUM  (i.  e.  leaf- 
branch)  ,  or  in  shorter  English  CLADOPHYLL. 

129.    Frondose   Stems.      Finally,  in  some   few 
124          phsenogamous  plants,  the  whole  vegetation  is  re- 
duced to  a  simple  leaf-like  expansion,  as  in  Duckweed  (Lemna) , 

FIG.  122.  Myrsiphyllum,  with  cladophylls  serving  for  foliage;  the  true  leaves  con- 
sisting of  minute  and  very  inconspicuous  scales  subtending  the  former. 

FIG.  123.  A  single  cladophyll  of  Ruscus  aculeatus  in  the  axil  of  a  scale-leaf,  bearing 
another  scale-leaf  on  the  middle  of  its  face,  and  flowers  in  the  axil  of  this. 

FIG.  124.    Lemna  minor,  a  common  Duckweed,  whole  plant  in  flower,  magnified. 


INTERNAL   STRUCTURE.  67 

Fig.  124.  Here  is  no  differentiation  whatever  into  stem  and 
foliage ;  but  the  expanded  floating  body  which  serves  for  both 
must  be  counted  as  stem  developed  horizontally  into  a  flat  plate, 
for  it  produces  a  root  from  the  under  surface  and  a  flower  from 
the  edge.  This  simplification  is  common  in  some  orders  of 
Cryptogamous  plants  ;  and  such  a  body,  which  answers  both  for 
stem  and  foliage,  is  termed  a  FROND,  from  the  Latin  frons,  which 
means  either  leaf  or  leafy  bough.  In  some  species  of  Lemna 
the  frond  is  thickened  or  plano-convex :  in  Wolffia,  the  simplest 
and  smallest  of  phaenogamous  plants,  it  is  a  globular  green  mass, 
seldom  much  larger  than  the  head  of  a  pin,  wholly  destitute  of 
root,  propagated  by  proliferous  budding  from  one  side,  and  from 
within  the  top  producing  a  flower  or  pair  of  flowers. 


§  3.  INTERNAL  STRUCTURE. 

130.  The  investigation  of  the  intimate  structure  of  the  stem, 
as  of  the  other  organs,  belongs  to  vegetable  anatomy  or  histology 
(treated  in  Part  II.)  ;  but  the  general  outlines  of  structure,  so 
far  as  is  requisite  to  the  explanation  of  what  is  visible  to  the 
naked  eye,  should  be  here  explained. 

131.  The  stems  of  phaenogamous  plants  anatomically  consist 
of  two  general  elements,  the  cellular  and  the  woody  ;  the  former 
exemplified  in  the  commoner  stems  by  the  pith  and  outer  bark,  the 
latter  by  the  wood.    Both  are  equally  composed  of  cells,  or  origi- 


nate  as  such  ;  but  those  which  form  the  woody  system  of  the  stem 
mainly  undergo,  at  a  very  early  period,  transformation  into  tubes, 
some  of  which  are  of  such  small  calibre  that  their  common  name 
of  fibres  is  not  inappropriate ;  others,  of  larger  size  or  ampler 
calibre,  take  the  name  of  ducts  or  vessels.  The  latter  are  almost 

FIG.  124°.  A  magnified  slice  of  a  portion  of  the  flower-stalk  of  Richardia  ^thiopica 
(the  so-called  Calla  Lily),  transverse  with  some  longitudinal  view :  mainly  parenchyma, 
the  cells  built  up  so  as  to  leave  comparatively  large  vacancies  (intercellular  spaces  or 
air-passages);  near  the  centre  a  cross-section  of  a  flbro- vascular  bundle,  and  next  the 
margin  or  rind  some  finer  ones. 


68  MOKPHOLOGY   OF   STEMS. 

always  associated  with  the  wood-cells,  so  that  they  are  in  a  general 
way  taken  together  as  constituting  the  wood,  or  woody  tissue,  and 
as  forming  what  is  more  definitely  termed  fibro-vascular  tissue  or, 
when  distinguishable  into  threads,  fibro-vascular  bundles.  These 
run  lengthwise  through  the  stem,  sometimes  as  such  separate 
threads,  sometimes  confluent  into  a  compact  structure.  The 
softer  or  at  least  the  non-fibrous  portions,  formed  of  comparatively 


short  and  commonly  thin- walled  cells,  form  cellular  tissue.  Its 
ordinary  form  (of  roundish,  cubical,  or  polyhedral  and  thin-walled 
cells)  is  called  parenchyma.  This  abounds  in  herbaceous  stems  or 
herbaceous  parts  :  in  trees  and  shrubs,  wood}'  tissue  largel}'  pre- 
vails ;  in  most  herbs,  it  forms  a  notable  portion ;  in  some  (especially 

FIG.  125.  Fibro-vascular  elements,  a.  Bast-cells  (long  wood-cells)  of  fibrous  bark 
of  Linden  or  Bass-wood,  b.  Some  wood-cells  and  (below)  a  duct,  and  c.  a  detached 
wood-cell  of  the  wood  of  same  tree,  equally  magnified  with  a.  d.  A  detached  wood- 
cell  from  a  shaving  of  White  Pine,  showing  the  peculiar  disk-like  markings,  e.  Portion 
of  same  shaving.  /.  Portion  of  a  dotted  duct  from  the  Vine,  evidently  made  up  of  a 
series  of  short  cells,  g.  Part  of  a  smaller  dotted  duct,  showing  no  appearance  of  such 
composition,  h,  i.  Spiral  ducts  or  vessels,  of  the  ordinary  kind.  j.  Spiral  duct  of 
Banana,  k.  Duct  from  Celery,  the  thread  within  spiral  or  annular  below,  reticulated 
above,  and  higher  passing  into  the  state  of  dotted  duct.  /.  Duct  from  Impatiens.  with 
the  open  spiral  passing  into  rings  at  the  middle.  All  magnified  somewhat  equally. 


INTERNAL   STRUCTURE.  69 

in  certain  aquatic  herbs),  it  is  reduced  to  a  few  threads  or 
vessels,  generally  delicate,  and  sometimes  obscure.  The  ac- 
companying anatomical  illustrations  (Fig.  124°,  125,  with  their 
explanations)  will  give  a  general  idea  of  the  nature  of  the  ana- 
tomical elements  of  the  stem. 

132.  In  the  forming  state,  the  whole  stem  is  parenchyma ;  but 
an  early  differentiation  takes  place,  converting  certain  portions 
into  woody  or  nbro-vascular  tissue.     This  is  arranged  in  two 
ways,  giving  rise  to  two  kinds  of  stem  in  phaenogamous  plants, 
which  have  been  termed  the  Endogenous  and  the  Exogenous,1 
meaning  inside  and  outside  growers. 

133.  The  two  plans  of  stem  are  usually  manifested  in  external 
conformation  as  well  as  internal  structure,  and  are  correlated 
with  important  differences  in  embryo,  foliage,  and  flower.2  Palms, 
Lilies,  Rushes,  and  Grasses  are  examples  of  the  endogenous 
class ;  the  ordinary  trees  and  shrubs,  especially  those  of  cool 
climates,  and  a  large  part  of  the  herbs,  are  of  the  exogenous  class. 
In  an  exogenous  stem,  the  wood  occupies  annual  concentric  layers, 
one  of  each  year's  growth ;   the  centre  is  occupied  by  a  pith, 
composed  of  parenchyma  only,  the  circumference  by  a  separable 
bark ;  so  that  a  cross-section  presents  a 

series  of  rings  or  circles  of  wood,  or  in 
the  first  year  one  ring,  surrounding  the 
pith  and  surrounded  by  the  bark.  An 
endogenous  stem  has  the  wood  in  distinct 
threads  or  fibro- vascular  bundles,  travers- 
ing the  cellular  system  or  parenchyma  with 
little  or  no  obvious  order,  and  presenting  126 

on  the  cross-section  the  divided  ends  of  these  bundles  in  the 
form  of  dots ;  these  usually  (but  not  always)  diffused  over 

1  Terms  introduced  by  DeCandolle,  following  the  ideas  of  Desfontaines, 
and  which  have  played  an  important  part  in  structural  and  systematic 
botany  ever  since  DeCandolle  adopted  these  names  as  those  of  the  two 
primary  divisions  of  phaenogamous  plants,  Exogence  and  Endogence.    But  it 
has  long  been  seen  that  the  name  of  the  second  kind  is  not  appropriate ;  and 
the  older  and  better  (though  longer)  names  of  Jussieu,  Monocotyledones  and 
Dicotyledones,  are  reverted  to.     Yet  the  Candollean  names  are  still  much 
employed,  with  due  explanation,  to  designate  the  two  kinds  of  structure  of 
the  stem. 

2  Yet  with  some  more  or  less  valid  exceptions,  as  when  the  annual  stem 
of  Podophyllum  and  the  rhizoma  of  Nymphaea,  among  dicotyledonous  plants, 
imitate  the  endogenous  structure ;  or  where  the  pith  of  an  evidently  exogenous 
stem,  as  in  the  Piperaceae,  has  scattered  woody  bundles  in  an  endogenous 
fashion ;  or  where  monocotyledonous  plants  have  all  their  woody  bundles  in 
a  definite  circle,  as  in  Luzula,  Croomia,  &c. 

FIG.  126.    Section  of  a  small  Palm-stem,  in  two  directions. 


70  MORPHOLOGY   OF   STEMS. 

the  whole  section,  or  when  few  in  number  of  somewhat  definite 
position  or  arrangement.  The  ordinary  appearance  of  such  a 
stem,  both  on  the  longitudinal  and  the  cross-section,  is  shown 
in  Fig.  126  ;  it  may  also  be  examined  in  the  Cane  or  Rattan, 
the  Bamboo,  and  in  the  annual  stalk  of  Indian  Corn  or  of 
Asparagus.  The  appearance  of  ordinary  wood  is  very  familiar. 

135.  The  newer  woody  bundles  of  an  endogenous  stem  are  vari- 
ously intermingled  with  the  old.  When  DeCandolle  gave  the  name, 
it  was  supposed,  from  Desfontaines's  researches,  that  the  older 
bundles  occupied,  or  came  at  length  to  occupy,  the  circumference 
of  the  trunk,  while  only  new  ones  were  formed  in  the  centre ; 
and  that  increase  in  diameter,  when  it  took  place  at  all,  resulted 
from  the  gradual  growth  and  distention  of  the  whole.     Hence 
the  contrasting  name  of  endogenous,  or  inside  growing,  and  for 
such  plants  the  name  of  ENDOGENOUS  PLANTS,  or  ENDOGENS. 
Our  actual  knowledge  of  the  structure  and  growth  of  these  stems, 
as  will  be  seen,  cannot  be  harmonized  with  this  view  in  any 
way  which  gives  to  the  name  endogenous  an  appropriate  signifi- 
cation.    The  name  continues  as  a  counterpart  to  the  more  correct 
one  of  exogenous,  and  as  a  survival  of  former  ideas. 

136.  The  Endogenous  Structure  (so  called)  of  the  stem  is  cor- 
related with  a  monocotjledonous  embryo  (39) ,  usually  with  a 
ternary  arrangement  in  the  flower  (322),  and  commonly  with 
parallel- veined  leaves.  (173.)     Endogens,  although   they  have 
many  herbaceous  and  a   few  somewhat  woody  representatives 
in  cool  temperate  climes,  mostly  attain  their  full  variety  of  fea- 
tures  and  rise   to   noble   arborescent   forms   under   a   tropical 
sun.     Yet  Palms  —  the  arboreous  type  of  the  class  —  do  extend 
as  far  north  in  this  country  as  the  coast  of  North  Carolina  (the 
natural  limit  of  the  Palmetto,  Fig.  126")  ;  while  in  Europe  the 
Date  and  the  Chamaerops  thrive  in  the  warmest  parts  of  the  Euro- 
pean shore  of  the  Mediterranean.     The  manner  of  their  growth 
gives  them  a  striking  appearance  ;  their  trunks  being  unbranched 
cylindrical  columns,  rising  to  the  height  of  from  thirty  to  one 
hundred  and  fifty  feet,  and  crowned  at  the  summit  with  a  simple 
cluster   of  peculiar  foliage.     Palms   generally  grow   from   the 
terminal  bud  alone,  and  perish  if  this  bud  be  destroyed ;  they 
grow  slowly,  and  bear  their  foliage  in  a  cluster  at  the  summit  of 
the  trunk,  which  consequently  forms  a  simple  cylindrical  column. 
But  in  some  instances  two  or  more  buds  develop,  and  the  stem 
branches,  rarely  and  accidentally  in  ordinary  species,  regularly 
in  the  Doum  Palm  of  Upper  Egypt,  and  in  the  Pandanus,  or 
Screw-Pine  (Fig.  69) ,  which  belongs  to  a  family  allied  to  Palms  : 
in  such  cases  the  branches  are  cylindrical.     But  when  lateraJ 


INTERNAL   STRUCTURE.  71 

buds  are  freely  developed  (as  in  the  Asparagus) ,  or  the  leaves 
are  scattered  along  the  stem  or  branches  by  the  full  development 
of  internodes  (as  in 
the  Bamboo,  Maize, 
&c. ) ,  they  gradually 
taper  upward  in  the 
manner  of  most  ex- 
ogenous stems. 

137.    This    kind 
of   stem    comprises 
several  subordinate 
types  as  to  internal 
structure,  which  to 
be  well  understood 
must  be  studied  his- 
tologically,  under  the 
microscope.1  To  one 
of  these,  by  no  means 
the  simplest,  belongs 
the   ordinary  palm- 
stem,   the   anatomy 
of  which  was  made 
classical   by   Mohl, 
and  has  been 
supplemented 
by      Nsegeli. 
In  this  a  large 
part    of    the 
bundles,  or  all 
of   the   more 

conspicuous  kind,  starting  from  the  base  of  the  leaf  to  which 
they  respectively  belong,  curve  inward  more  or  less  strongly 
toward  the  centre  of  the  stem,  and  thence  gradually  outward 
as  they  descend  until  they  reach  the  rind,  in  which  the 
attenuated  lower  extremity  mostly  terminates.  Consequently,  the 
bundles  from  different  heights  cross  in  their  course,  somewhat 

1  For  the  best  and  most  accessible  memoir  on  the  subject,  of  recent  date, 
see  Guillaud,  Recherches  sur  1'Anatomie  comparee  et  le  Developpement  des 
Tissus  de  la  Tige  dans  les  Monocotyle'dones,  published  in  Ann.  Sci.  Nat. 
ser.  6,  v.  1-176,  1877.  Six  types  of  the  stem  of  Monocotyledons  are  here 
recognized  by  anatomical  characters  and  modes  of  growth,  one  of  them 
having  four  modifications. 

FIG.  126"  Sabal  Palmetto  in  various  stages;  also  the  Yucca  aloifolia  or  Spanish 
Bayonet. 


72  MORPHOLOGY   OF   STEMS. 

as  shown  in  Fig.  127.  It  is  partly  owing  to  this  connection  of 
these  fibres  with  the  rind  that  the  latter  is  not  separable  from 
the  stem.  In  some  Palms,  and  in  Grasses, 
there  is  no  marked  distinction  between 
the  wood  and  rind,  or  no  proper  rind  at 
all.  In  others,  such  as  the  Palmetto 
(Fig.  126),  there  is  a  marked  rind 
or  false  bark,  which  receives  independent 
fibro- vascular  bundles  from  the  leaf-stalks, 
and  is  traversed  by  them  in  parallel  lines. 
In  Grass-stems,  and  others  with  long  inter- 
nodes  and  closed  nodes,  the  fibro-vascular 
bundles  all  run  approximately  straight 
and  parallel  through  the  internodes,  but  are 
intricate  and  anastomosed  in  the  nodes. 
The  whole  centre  of  the  internodes,  when 

hot  hollow  or  before  it  becomes  so,  is  occupied  b}'  a  true  pith, 
(ike  that  of  an  Exogen,  and  in  some  cases  equally  destitute  of 
fibro-vascular  bundles,  but  often  with  scattered  ones,  after  the 
manner  of  certain  Exogens  anomalous  in  this  respect,  such  as 
Nj-ctaginaceae  and  some  Araliaceae.  Endogenous  stems  of 
simpler  structure,  as  in  herbaceous  Liliacese,  Commelynacese, 
&c.,  have  a  distinct  cortical  portion  (at  least  in  the  root-stock 
or  portion  of  stem  properly  comparable  with  palm-trunks  and 
the  like)  ;  but  this  is  mostly  destitute  of  fibro-vascular  bundles. 
Most  of  them  have  two  kinds  of  vascular  bundles,  one 
of  which  not  rarely  occupies  an  exact  circle  in  the  line  of 
division  between  the  cortical  and  medullary  portion  (between 
bark  and  pith),  and  the  other  is  within  this  circle,  either  of 
very  few  and  scattered  bundles,  as  in  Convallaria  majalis,  or 
numerous  and  scattered,  as  in  Uvularia  and  the  leafy  stems  of 
Tradescantia  Virginica  ;  or  these  bundles  are  few  and  arranged 
nearly  in  an  inner  circle  close  around  the  centre.  Finally,  Luzula 
and  Croomia  have  only  one  kind  of  bundles,  answering  to  the 
outer  ones  of  Convallaria ;  in  other  words,  the  woody  system 
forms  a  simple  circle,  dividing  a  purely  cellular  medullary  from  a 
similar  cortical  portion,  thus  closely  imitating  an  herbaceous 
exogenous  stem  of  the  same  age. 

138.  An  annual  endogenous  stem  increases  in  diameter  by 
general  growth  until  it  attains  its  limit.  Ligneous  and  enduring 
stems  increase  similarly  up  to  a  certain  period.  Then  the  rind 


FIG.  127.    Diagrammatic  view  of  the  curved  course  of  the  flbro-vascular  bundles 
ill  a  palm-trunk. 


INTERNAL   STRUCTURE. 


73 


sooner  or  later  ceases  to  distend  or  adapt  itself  to  further  in- 
crease in  diameter,  and  there  is  no  interior  provision  for  indefinite 
increase  in  the  greater  number  of  woody  endogenous  trunks.  But 
in  Dracaena  (Dragon-trees) ,  in  the  arborescent  Yuccas,  and  the 
like,  the  zone  intermediate  between  the  cortical  and  interior  re- 
gion, which  is  for  a  time  active  in  many  Endogens,  here  grows 
continuously  and  indefinitely.  Such  trunks  increase  in  diameter 
throughout  life ;  they  may  attain  a  very  great  age  (as  some 
Dragon-trees  have  done)  ;  and  they  imitate  exogenous  trunks 
to  a  considerable  extent  in  mode  of  growth. 

139.  The  wood  of  an  endogenous  woody  stem  is  hardest  and 
most  compact  at  the  circumference ;  in  palm-stems  commonly 
it  is  largely  mixed  with  parenchyma  or  pith  at  the  centre,  even 
in  old  trunks. 

140.  The  Exogenous  Structure,  that  of  ordinary  wood,  is  char- 
acterized by  the  formation  of  a  distinct  zone  of  wood  between  a 
central  cellular  medullary  portion   (pith)   and  an  outer  chiefly 
cellular  portion  (bark) ,  traversed  by  plates  from  the  pith  (medul- 
lary rays) ,  and  by  increasing  from  the  outer  surface  of  this  zone 
between  wood  and  bark,  the  increase  in  enduring  stems  consist- 
ing of  definite  concentric  annual  layers. 

141.  Its  Beginning,  at  the  earliest  growth  of  the  embryo,  is  in 
the  appearance  of  a  few  ducts  (Fig.  125,/-/),  at  definite  points 
in  the  common  parenchyma  of  the  initial  stem  (four  equidistant 
ones  in  the  Sugar  Maple)  ;  each  is  soon  surrounded  by  incipient 


proper  wood-cells  (Fig.  125, 5,  c),  together  forming  a  fibro-vascular 
bundle  or  thread.  Additional  ones  are  intercalated  as  the  second 
and  third  internodes  develop,  and  so  a  column  (in  cross- 
section  a  ring)  of  wood  is  produced,  always  so  arranged  as  to 


FIG.  128.  Diagram  of  a  cross-section  of  a  forming  seedling  stem,  showing  the 
manner  in  which  the  young  wood  is  arranged  in  the  cellular  system. 

FIG.  129.  The  same  at  a  later  period,  the  woody  bundles  increased  so  as  nearly  to 
fill  the  circle. 

FFG.  130.  The  same  at  the  close  of  the  season,  where  the  wood  has  formed  a  com- 
plete circle,  interrupted  only  by  the  medullary  rays,  which  radiate  from  the  pith  to 
the  bark. 


74 


MORPHOLOGY  OF   STEMS. 


JO* 


surround  a  purely  cellular  central  part  (the  pith),  while  sur- 
rounded by  a  cellular  external  rind,  the  bark,  or  outer  bark. 
The  diagrams  (Fig.  128-130)  rudely  show  some  stages  in  the 
formation  of  the  zone  of  wood.  The  fibro- vascular  bundles 
originate  in  the  bases  of  the  leaves,  and  develop  outward  into  the 
forming  leaves  as  well  as  downward  into  the  forming  stem. 

142.  First  Year's  Growth.  The  wood,  even  in  a  herbaceous 
or  annual  stem,  at  the  completion  of  the  first  year's  growth, 

forms  a  zone  or  tube, 
enclosing  the  pith.  But 
it  is  traversed  by  plates 
(in  cross-section  lines) 
of  parenchyma,  or  eel 
lular  tissue  of  the  same 
nature  as  the  pith, 
which  radiate  from 
that  to  the  bark,  and 
thus  divide  the  wood 
into  wedges.  These 
lines,  forming  what  is 
called  the  silver-grain 
in  wood,  are  the  MED- 
ULLARY RAYS.  They 
represent  the  cellular 
system  of  the  wood  it- 
self, or  untransformed 
parenchyma.  Being 
pressed  by  the  woody 
wedges,  their  cells  are 
laterally  flattened.  In 
some  stems,  the  med- 
ullary rays,  or  many 
of  them,  are  comparatively  broad  and  conspicuous;  in  others, 
thin  and  inconspicuous  or  irregular.  The  growth  of  the  woody 
wedges  is  soon  complete,  except  at  the  outer  portion,  next 
the  bark :  here  they  usually  continue  to  grow  through  the 
season ;  that,  is  the  wood  grows  externally.  The  general  ana- 

FIG.  131.  Longitudinal  and  transverse  section  of  a  stem  of  the  Soft  Maple  (Acer 
dasycarpum ),  at  the  close  of  the  first  year's  growth  ;  of  the  natural  size. 

FIG.  132.  Portion  of  the  same,  magnified,  showing  the  cellular  pith,  surrounded  by 
the  wood,  and  that  by  the  bark. 

FIG.  133.  More  magnified  slice  of  the  same,  reaching  from  the  bark  to  the  pith : 
a.  part  of  the  pith ;  b.  vessels  of  the  medullary  sheath ;  c.  the  wood ;  d ,  d.  dotted  ducts  in 
the  wood ;  e,  e.  annular  ducts ;  /.  the  liber,  or  inner  fibrous  bark ;  g.  the  cellular  envelope, 
or  green  bark ;  h.  the  corky  envelope ;  t.  the  skin  or  epidermis ;  k.  one  of  the  medullary 
raya,  seen  on  the  transverse  section. 


INTERNAL   STRUCTURE.  75 

tomical  structure  of  a  woody  exogenous  stem  of  a  year  old  is 
displayed  in  the  Fig.  131-133.  Viewing  the  parts  particularly, 
and  in  order  from  centre  to  circumference,  there  is,  — 

1st.  The  Pith  or  Medulla,  consisting  entirely  of  soft  and  rather 
large  thin- walled  cells,1  gorged  with  sap  or  other  nourishing 
matter  during  the  growing  state,  becoming  light,  dry,  and  empty 
when  effete. 

2nd.  The  Layer  of  Wood,  traversed  by  the  medullary  rays.  In 
Pines  and  other  Coniferae,  the  wood  is  of  uniform  structure,  being 
wholly  composed  of  a  woody  tissue  with  peculiar  markings  (Fig. 
125,  d,  e)  :  in  other  wood,  ducts  of  one  or  more  sorts  occur ;  the 
most  conspicuous  being  what  are  termed  dotted  ducts.  These 
are  so  large  as  to  be  evident  to  the  naked  eye  in  many  ordi- 
nary kinds  of  wood,  especially  where  they  are  accumulated  in 
the  inner  portion  of  the  layer,  as  in  the  Chestnut  and  Oak.  In 
the  Maple,  Plane,  &c.,  they  are  rather  equably  scattered  through 
the  annual  layer,  and  are  too  small  to  be  seen  by  the  naked  eye. 
Next  the  pith,  i.  e.  in  the  very  earliest  formed  part  of  the  wood, 
some  spiral  ducts  are  uniformly  found,  and  this  is  the  only  part 
of  the  exogenous  stem  in  which  these  ordinarily  occur.  They 
may  be  detected  by  breaking  a  woody  twig  in  two,  after  dividing 
the  bark  and  most  of  the  wood  by  a  circular  incision,  and  then 
pulling  the  ends  gently  asunder,  when  their  spirally  coiled  fibres 
are  readily  drawn  out  as  gossamer  threads.  As  these  spiral 
ducts  form  a  circle  immediately  surrounding  the  pith,  they  have 
collectively  been  termed  the  medullary  sheath,  but  they  hardly 
deserve  a  special  name.  The  vertical  section  in  Fig.  133  divides 
one  of  the  woody  wedges,  and  shows  no  medullary  ray  ;  but  there 
is  one  at  the  posterior  edge  of  the  transverse  section.  But,  in  the 
much  more  diagramatic  Fig.  134,  the  section  is  made  so  as  to  show 
the  surface  of  one  of  these  plates,  or  medullary  rays,  passing  hori- 
zontally across  it,  connecting  the  pith  (jo)  with  the  bark  (b) . 
These  medullary  rays  form  the  silver-grain  (as  it  is  termed) ,  which 
is  so  conspicuous  in  the  Maple,  Oak,  &c.,  and  which  gives  the 
glimmering  lustre  to  many  kinds  of  wood  when  cut  in  this  direc- 
tion. A  section  made  as  a  tangent  to  the  circumference,  and 
therefore  perpendicular  to  the  medullary  rays,  brings  their  ends 
to  view,  as  in  Fig.  135,  much  as  they  appear  on  the  surface  of  a 
piece  of  wood  from  which  the  bark  is  stripped.  They  are  here 
seen  to  be  composed  of  parenchyma,  and  to  represent  the  horizon- 

1  In  rare  instances,  a  few  fibro-vascular  threads  are  found  dispersed 
through  the  pith,  presenting  a  somewhat  remarkable  anomaly.  This 
occurs  in  Aralia  racemosa,  and  more  strikingly  in  Mirabilis  and  other 
Nyctaginaceae,  and  in  Piperaceae.  (133.  foot-note.) 


76 


MORPHOLOGY   OF   STEMS. 


tal  system  of  the  wood,  or  the  woof,  into  which  the  vertical  woody 
fibre,  &c.,  or  warp,  is  interwoven.  The  inspection  of  a  piece  of  oak 
or  maple  wood  at  once  shows  the  pertinency  of  this  illustration. 


3rd.  The  Bark  or  rind.  This  at  first  consisted  of  simple 
parenchyma,  like  that  of  the  pith,  except  for  the  green  color 
developed  in  it,  the  same  as  that  which  gives  verdancy  to  foliage. 
This  green  matter  is  formed  in  the  cells  of  ah1  such  parts  when 
exposed  to  light,  consists  of  green  grains  of  somewhat  complex 
chemical  composition,  has  important  functions  to  perform  in 
assimilation  (i.  e.  in  the  conversion  of  the  plant's  crude  food  into 
vegetable  matter)  ,  and  is  named  CHLOROPHYLL,  i.  e.  leaf-green. 
The  completed  bark,  when  all  its  parts  are  apparent,  as  espe- 
cially in  most  trees  and  shrubs,  is  composed  of  three  strata,  of 
which  the  green  bark,  the  most  conspicuous  in  the  young  shoot, 
is  the  middle  layer,  therefore  named  the  MESOPHLCEUM.  This  is 
soon  covered,  and  the  green  color  obscured,  by  a  superficial 
stratum  of  cells,  generally  of  some  shade  of  ash-color  or  brown, 
occasionally  of  brighter  tints,  which  gives  to  the  twigs  of  trees 
and  shrubs  the  hue  characteristic  of  each  species,  the  CORKY 
ENVELOPE  or  layer,  or  EPIPHLGEUM.  The  latter  name  denotes  its 
external  position  ;  the  former,  that  it  is  the  la}rer  which,  when 
much  developed,  forms  the  cork  of  Cork-Oak  and  those  corky 
expansions  which  are  so  conspicuous  on  the  twigs  of  the  Sweet 
Gum  (Liquidambar)  ,  and  on  some  of  our  Elms  (Ulmus  alata 
and  racemosa)  .  It  also  forms  the  paper-h'ke  exfoliating  layers 
of  Birch-bark.  It  is  composed  of  laterally  flattened  parenchy- 
matous  cells,  much  like  those  of  the  EPIDERMIS  (Fig.  133,  «), 
which  directly  overlies  it,  and  forms  the  skin  or  surface  of  the 


FIG.  134.  Vertical  section  through  the  wood  of  a  branch  of  the  Maple,  a  year  old, 
so  as  to  show  one  of  the  medullary  rays,  passing  transversely  from  the  pith  (p)  to  the 
bark  (6):  magnified.  But  a  section  can  seldom  be  made  so  as  to  show  one  unbroken 
plate  stretching  across  the  wood,  as  In  this  instance. 

FIG.  136.    A  vertical  section  across  the  ends  of  the  medullary  ray^:  magnified. 


INTERNAL  STRUCTURE. 


77 


stem,  and  of  the  whole  plant.  Lastly,  the  inner  bark,  accord- 
ingly named  ENDOPHLCEUM,  takes  the  special  name  of  LIBER,  and 
is  the  most  important  portion  of  the  bark  in  the  stems  of  trees 
and  shrubs.  Complete  and  well-developed  liber,  like  that  of 
Linden  or  Basswood,  contains  two  peculiar  kinds  of  cells  in 


addition  to  common  parenchyma,  both  of  the  fibrous  or  vascular 
class:  viz.,  1.  CRIBRIFORM  or  SIEVE-CELLS,  a  sort  of  ducts  the 
walls  of  which  have  open  slits,  through  which  they  communicate 
with  each  other ;  2.  BAST  or  BAST-CELLS,  the  fibre-like  cells 
which  give  to  the  kinds  of  inner  bark  that  largely  contain  them 

FIG.  135°.  Portion  of  a  transverse  section  (above),  and  a  corresponding  vertical  sec- 
tion (below),  magnified,  reaching  from  the  pith  (p)  to  the  epidermis  (e)  of  a  stem  of 
Negundo,  a  year  old :  B.  the  bark ;  W.  the  wood ;  and  C.  the  cambium-layer,  as  found 
in  February.  The  parts  referred  to  by  small  letters  are :  p.  a  portion  of  the  pith : 
mr.  small  portion  of  a  medullary  ray  where  it  runs  into  the  pith;  four  complete  med- 
ullary rays  as  seen  on  a  transverse  section,  appear  in  the  upper  figure,  running  from 
pith  to  bark :  ms,  medullary  sheath,  a  circle  of  spiral  unreliable  ducts,  one  seen  length- 
wise with  uncoiling  extremity  in  the  lower  figure :  w,  w.  woody  tissue :  dd.  one  of  the 
dotted  ducts  interspersed  in  the  wood :  cl.  cambium-layer  or  zone  of  new  growth  of 
wood  and  inner  bark :  l-b.  liber  or  inner  bark,  the  inner  portion  of  which  is  here  cellular, 
the  outer  (6)  composed  of  slender  and  thick-walled  bast-cells  or  true  liber-cells: 
ge.  green  envelope  or  inner  cellular  bark :  ce.  corky  envelope  or  outer  cellular  bark : 
e.  epidermis. 


78  MORPHOLOGY   OF   STEMS. 

their  strength  and  toughness.  They  are  like  wood-cells  except 
in  their  greater  length  and  flexibility,  and  in  the  thickness  of 
their  walls,  which  greatly  exceeds  the  calibre.  This  is  the 
material  which  gives  to  the  bast  or  inner  bark  of  Basswood,  &c., 
the  strength  and  pliability  that  adapts  it  for  cordage  and  for 
making  mats :  it  is  the  material  of  linen,  and  the  like  textile 
fibres.  (For  a  view  of  the  whole  composition  and  structure  of  a 
woody  stem  at  the  close  of  the  first  year's  growth,  and  immedi- 
ately before  that  of  the  second  year  begins,  see  Fig.  135a.) 

143.  Annual  Increase  in  Diameter.  An  herbaceous  stem  does 
not  essentially  differ  from  a  woody  one  of  the  same  age,  except 
that  the  wood  forms  a  less  compact  or  thinner  zone ;  and  the 
whole  perishes,  at  least  down  to  the  ground,  at  the  close  of  the 
season.  But  a  woody  stem  makes  provision  for  continuing  its 
growth  from  year  to  year.  As  the  layer  of  wood  continues  to 
increase  in  thickness  throughout  the  season,  by  the  multiplication 
of  cells  on  its  outer  surface,  between  it  and  the  bark,  and  when 
growth  ceases  this  process  of  cell-multiplication  is  merely  sus- 
pended, so  there  is  always  a  zone  of  delicate  young  cells  in- 
terposed between  the  wood  and  the  bark.  This  is  called  the 
CAMBIUM,  or,  better,  the  CAMBIUM-LAYER.  It  is  charged  with 
organizable  matter,  which  is  particularly  abundant  and  mucila- 
ginous in  spring  when  growth  recommences.  This  mucilaginous 
matter  was  named  Cambium  by  the  older  botanists  :  the}7  sup- 
posed—  as  is  still  popularly  thought  —  that  the  bark,  then  so 
readily  separable,  really  separated  from  the  wood  in  spring,  that  a 
quantity  of  rich  mucilaginous  sap  was  poured  out  between  them 
and  became  organized  into  a  tissue,  the  inner  part  becoming  new 
wood,  the  outer,  new  bark.  But  delicate  slices  show  that  there 
is  then  no  more  interruption  of  the  wood  and  inner  bark  than 
at  any  other  season.  The  bark,  indeed,  is  then  very  readily 
detached  from  the  wood,  because  the  cambium-layer  is  gorged 
with  sap ;  but  such  separation  is  effected  by  the  rending  of  a 
delicate  forming  tissue.  And  if  some  of  this  apparent  mucilage 
be  scraped  off  from  the  surface  of  the  wood,  and  examined  under 
a  good  microscope,  it  will  be  seen  to  be  a  thin  stratum  of  young 
wood-cells,  with  the  ends  of  medullary  rays  here  and  there  in- 
terspersed. The  inner  portion  of  the  cambium-layer  is  therefore 
nascent  wood,  and  the  outer  is  nascent  bark.  As  the  cells  of 
this  layer  multiply,  the  greater  number  lengthen  vertically  into 
woody  tissue :  some  are  transformed  into  ducts  ;  and  others, 
remaining  as  parenchyma,  continue  the  medullary  rays  or  com- 
mence new  ones.  In  this  way,  a  second  la}-er  of  wood  is  formed 
the  second  season  over  the  whole  surface  of  the  former  layer 


INTERNAL  STRUCTURE.  79 

between  it  and  the  bark  ;  and  this  is  continuous  with  the  woody 
layer  of  the  new  roots  below  and  of  the  leafy  shoots  of  the  sea- 
son above.  Each  succeeding  year  another  layer  is  added  to  the 
wood  in  the  same  manner,  coincident  with  the  growth  in  length 
by  the  development  of  the  buds.  A  cross-section  of  an  exoge- 
nous stem,  therefore,  exhibits  the  wood  disposed  in  concentric 
rings  between  the  bark  and  the  pith ;  the  oldest  lying  next  the 
latter,  and  the  youngest  occupying  the  circumference.  Each 
layer  being  the  product  of  a  single  year's  growth,  the  age  of  an 
exogenous  tree  may,  in  general,  be  correctly  ascertained  by 
counting  the  rings  in  a  cross-section  of  the  trunk.1 

144.  Demarcation  of  the  Annual  Layers  results  from  two  or  more 
causes,  separate  or  combined.  In  oak  and  chestnut  wood,  and 
the  like,  the  layers  are  strongly  denned  by  reason  of  the  accumu- 
lation of  the  large  dotted  ducts  (here  of  extreme  size  and  in 
great  abundance)  in  the  inner  portion  of  each  layer,  where  their 
open  mouths  on  the  cross-section  are  conspicuous  to  the  naked 
e}7e,  making  a  strong  contrast  between  the  inner  porous  and  the 
exterior  solid  part  of  the  successive  layers.  In  maple  and  beech 
wood,  however,  the  ducts  are  smaller,  and  are  dispersed  through- 
out the  whole  breadth  of  the  layer ;  and  in  coniferous  wood,  viz. 
that  of  Pine,  Cypress,  &c.,  there  are  no  ducts  at  all,  but  only  a 
uniform  woody  tissue  of  a  peculiar  sort.  In  all  these,  the  de- 
marcation between  two  layers  is  owing  to  the  greater  fineness  of 
the  wood-cells  formed  at  the  close  of  the  season,  viz.  those  at 
the  outer  border  of  the  layer,  while  the  next  layer  begins,  in  its 

1  The  annual  layers  are  most  distinct  in  trees  of  temperate  climates  like 
ours,  where  there  is  a  prolonged  period  of  total  repose,  from  the  winter's 
cold,  followed  by  a  vigorous  resumption  of  vegetation  in  spring.  In  tropical 
trees,  they  are  rarely  so  well  defined ;  but  even  in  these  there  is  generally  a 
more  or  less  marked  annual  suspension  of  vegetation,  occurring,  however, 
in  the  dry  and  hotter,  rather  than  in  the  cooler  season.  There  are  numerous 
cases,  moreover,  in  which  the  wood  forms  a  uniform  stratum,  whatever  be 
the  age  of  the  trunk,  as  in  the  arborescent  species  of  Cactus  ;  or  where  the 
layers  are  few  and  by  no  means  corresponding  with  the  age  of  the  trunk,  as 
in  the  Cycas. 

In  many  woody  climbing  or  twining  stems,  such  as  those  of  Clematis, 
Aristolochia  Sipho,  and  Menispermum  Canadense,  the  annual  layers  are 
rather  obscurely  marked,  while  the  medullary  rays  are  unusually  broad ; 
and  the  wood,  therefore,  forms  a  series  of  separable  wedges  disposed  in  a 
circle  around  the  pith.  In  the  stem  of  Bignonia  capreolata,  the  annual  rings, 
after  the  first  four  or  five,  are  interrupted  in  four  places,  and  here  as  many 
broad  plates  of  cellular  tissue,  belonging  properly  to  the  bark,  are  inter- 
posed, passing  at  right  angles  to  each  other  from  the  circumference  towards 
the  centre,  so  that  the  transverse  section  of  the  wood  nearly  resembles  a 
Maltese  cross.  But  these  are  exceptional  cases,  which  scarcely  require 
notice  in  a  general  view. 


80  MORPHOLOGY  OF   STEM*}. 

rigorous  vernal  growth,  with  much  larger  cells,  thus  marking 
an  abrupt  transition  from  one  layer  to  the  next.  Besides  being 
finer,  the  later  wood-cells  of  the  season  are  commonly  flattened 
antero-posteriorly,  probably  by  growing  under  greater  pressure. 

145.  Each  layer  of  wood,  once  formed,  remains  essentially 
unchanged  in  position  and  dimensions.     But,  in  trunks  of  con- 
siderable age,  the  older  layers  undergo  more  or  less  change  in 
color,  density,  perviousness  to  moisture,  &c. 

146.  Sap-wood    (ALBURNUM).      In   the   plantlet  and   in  the 
developing  bud,  the  sap  ascends  through  the  whole  tissue,  of 
whatever  sort :  at  first  through  the  parenchyma,  for  there  is  then 
no  other  tissue ;  and  the  transmission  is  continued  through  it, 
especially  through  its  central  portion,  or  the  pith,  in  the  growing 
apex  of  the  stem  throughout.    But,  in  the  older  parts  below,  the 
pith,  soon  drained  of  sap,  becomes  filled  with  air  in  its  place, 
and  thenceforth  it  bears  no  part  in  the  plant's  nourishment.     As 
soon  as  wood-cells  and  ducts  are  formed,  they  take  an  active 
part  in  the  conveyance  of  sap,  for  which  their  tubular  and  ca- 
pillary character  is  especially  adapted.     But,  the  ducts  in  older 
parts,  except  when  gorged  with  sap,  contain  air  alone ;  and  in 
woody  trunks  the  sap  continues  to  rise  year  after  year  to  the 
places  where  growth  is  going  on,  mainty  through  the  proper 
woody  tissue  of  the  wood.     In  this  transmission,  the  new  layers 
are  most  active  ;  and  these  are  in  direct  communication  with  the 
new  roots  on  the  one  hand  and  with  the  buds  or  shoots  and  leaves 
of  the  season  on  the  other.     So,  by  the  formation  of  new  annual 
layers  outside  of  them,  the  older  ones  are  each  year  removed  a 
step  farther  from  the  region  of  growth ;  or  rather  the  growing 
stratum,  which  connects  the  fresh  rootlets  that  imbibe  with  the 
foliage  that  elaborates  the  sap,  is  each  year  removed  farther  from 
them.     The  latter,  therefore,  after  a  few  years,  cease  to  convey 
sap,  as  they  have  long  before  ceased  to  take  part  in  any  vital 
operations.     The  cells  of  the  older  layers,  also,  usually  come  to 
have  thicker  walls  and  smaller  calibre  than  those  of  the  newer. 
Thus  arises  a  distinction  —  sometimes  obscurely  marked,  some- 
times abrupt  and  conspicuous  —  into  sap-wood  and  heart-wood. 
The  former  is  the  popular  name  given  to  the  outer  and  newer 
layers  of  softer,  more  open,  and  bibulous  wood.    The  early  physi- 
ologists named  it  alburnum  from  its  white  or  pale  color.     Being 
more  or  less  sappy,  or  containing  soluble  organic  matter,  and 
readily  imbibing  moisture,  this  part  of  the  wood  is  liable  to  decay, 
and  it  is  therefore  discarded  from  timber  used  for  construction. 

147.  Heart-wood  (or  DURAMEN,  so  called  from  its  greater  hard- 
ness or  durability)  is  the  older  and  mature  portion  of  the  wood, 


INTERNAL   STRUCTURE.  81 

In  all  trees  which  have  the  distinction  between  the  sap-wood  and 
heart- wood  well  marked,  the  latter  acquires  a  deeper  color,  and 
that  peculiar  to  the  species,  such  as  the  dark  brown  of  the  Black 
Walnut,  the  blacker  color  of  the  Ebony,  the  purplish-red  of  Red 
Cedar,  and  the  bright  yellow  of  the  Barberry.  These  colors  are 
owing  to  special  vegetable  products,  or  sometimes  to  alterations 
resulting  from  age.  In  the  Red  Cedar,  the  deep  color  belongs 
chiefly  to  the  medullary  rays.  In  many  of  the  softer  woods,  there 
is  little  change  in  color  of  the  heart- wood,  except  from  incipient 
decay,  as  in  the  White  Pine,  Poplar,  Tulip-tree,  &c.  The 
heart-wood  is  no  longer  in  any  sense  a  living  part :  it  may  perish, 
as  it  frequently  does,  without  affecting  the  life  or  health  of  the 
tree. 

148.  The  Growth  and  Duration  of  the  Bark,  also  the  differences 
in  structure,  are  much  more  various  than  of  the  wood.   Moreover, 
the  bark  is  necessarily  subject  to  grave  alterations  with  advanc- 
ing age,  on  account  of  its  external  position  ;  to  distention  from 
the  constantly  increasing  diameter  of  the  stem  within,  and  to 
abrasion  and  decay  from  the  influence  of  the  elements  without. 
It  is  never  entire,  therefore,  on  the  trunks  of  large  trees ;  but 
the  dead  exterior  parts,  no  longer  able  to  enlarge  with  the  en- 
larging wood,  are  gradually  fissured  and  torn,  and  crack  off  in 
strips  or  pieces,  or  disappear  by  slow  decay.     So  that  the  bark 
of  old  trunks  bears  only  a  small  proportion  in  thickness  to  the 
wood,  even  when  it  makes  an  equal  amount  of  annual  growth. 

149.  The  three  parts  of  the  bark   (142),  for  the  most  part 
readily  distinguishable  in  the  bark  of  young  shoots,  grow  inde- 
pendently, each  by  the  addition  of  new  cells  to  its  inner  face,  so 
long  as  it  grows  at  all.     The  green  layer  commonly  does  not 
increase  after  the  first  year ;  the  opaque  corky  layer  soon  excludes 
it  from  the  light ;  and  it  gradually  perishes,  never  to  be  renewed. 
The  corky  layer  usually  increases  for  a  few  years  only,  by  the 
formation  of  new  tabular  cells  :  occasionally  it  takes  a  remarkable 
development,  forming  the  substance  called   Cork,  as  in  the  Cork 
Oak,  and  the  thin  and  parchment-like  layers  of  the  White  and 
Paper  Birches. 

150.  The  liber,  or  inner  bark,  continues  its  growth  through- 
out the  h'fe  of  the  exogenous  tree,  by  an  annual  addition  from 
the  cambium-layer  applied  to  its  inner  surface.     Sometimes  this 
growth  is  plainly  distinguishable  into  layers,  corresponding  with 
or  more  numerous  than  the  annual  layers  of  the  wood :  often, 
there  is  scarcely  any  trace  of  such  layers  to  be  discerned.     In 
composition  and  appearance,  the  liber  varies  greatly  in  different 
plants,  especially  in  trees  and  shrubs.     That  of  Basswood  or 


82  MORPHOLOGY  OP   STEMS. 

Linden,  and  of  other  plants  with  a  similar  fibrous  bark,  may  be 
taken  as  best  representing  the  liber.  Here  it  consists  of  alter- 
nate strata  of  fibrous  bast,  and  of  the  peculiar  liber-cells  called 
sieve-cells,  in  which  nourishing  matter  is  especially  contained 
and  elaborated.  While  the  latter,  or  their  equivalents,  occur 
and  play  an  important  part  in  all  inner  bark,  the  bast-cells  are 
altogether  wanting  in  the  bark  of  some  plants,  and  are  not  pro- 
duced after  the  first  year  in  many  others.  The  latter  is  the  case 
in  Negundo,  where  abundant  bast-cells,  like  those  of  Basswood, 
compose  the  exterior  portion  of  the  first  year's  liber,  but  none 
whatever  are  formed  in  the  subsequent  layers.  In  Beeches  and 
Birches,  also,  a  few  bast-cells  are  produced  the  first  year,  but 
none  afterwards.  In  Maples,  a  few  are  formed  in  succeeding 
years.  In  the  Pear,  bast-cells  are  annually  formed,  but  in  very 
small  quantity,  compared  with  the  parenchymatous  part  of  the 
liber.  In  Pines,  at  least  in  White  Pines,  the  bark  is  nearly  as 
homogeneous  as  the  wood,  the  whole  liber,  except  what  answers 
to  the  medullary  rays,  consisting  of  one  kind  of  cells,  resembling 
those  of  bast  or  of  wood  in  form,  but  agreeing  with  the  proper 
liber-cells  in  their  structure  and  markings. 

151.  The  bark  on  old  stems  is  constantly  decaying  or  falling 
away  from  the  surface,  without  any  injury  to  the  tree ;  just  as 
the  heart-wood  within  may  equally  decay  without  harm,  except 
by  mechanically  impairing  the  strength  of  the  trunk.     There  are 
great  differences  as  to  the  time  and  manner  in  which  the  older 
bark  of  different  shrubs  and  trees  is  thrown  off.     Some  have 
their  trunks  invested  with  the   liber  of  many  years'  growth, 
although  only  the  innermost  layers  are  alive  ;  in  others,  it  scales 
off  much  earlier.     On  the  stems  of  the  common  Honeysuckle,  of 
the  Nine-Bark  (Spiraea  opulifolia) ,  and  of  Grape-vines  (except 
Vitis  vulpina),  the  liber  lives  only  one  season,  and  is  detached 
the  following  year,  hanging  loose  in  papery  layers  in  the  former 
species,  and  in  fibrous  shreds  in  the  latter. 

152.  While  the  newer  la}Ters  of  the  wood  abound  in  crude  sap, 
which  they  convey  to  the  leaves,  those  of  the  inner  bark  abound 
in  elaborated  sap,  which  they  receive  from  the  leaves  and  convey 
to  the  cambium-layer  or  zone  of  growth.     The  proper  juices  and 
peculiar  products  of  plants  are  accordingly  found  in  the  foliage 
and  the  bark,  especially  in  the  latter.     In  the  bark,  therefore 
(either  of  the  stem  or  of  the  root) ,  medicinal  and  other  principles 
are  usually  to  be  sought,  rather  than  in  the  wood.     Nevertheless, 
as  the  wood  is  kept  in  connection  with  the  bark  by  the  medullary 
rays,  many  products  which  probably  originate  in  the  former  are 
deposited  in  the  wood. 


INTERNAL  STRUCTURE.  83 

153.  The  Living  Parts  of  a  tree  or  shrub,  of  the  exogenous 
kind,  are  obviously  only  these:    1st,  The  summit  of  the  stem 
and  branches,  with  the  buds  which  continue  them  upwards  and 
annually  develop  the  foliage.     2d,  The  fresh  roots  and  rootlets 
annually  developed  at  the  opposite  extremity.     3d.  The  newest 
strata  of  wood  and  bark,  and  especially  the  interposed  cambium- 
layer,  which,  annually  renewed,  maintain  a  living  communication 
between  the  rootlets  on  the  one  hand  and  the  buds  and  foliage 
on  the  other,  however  distant  they  at  length  may  be.     These  are 
all  that  is  concerned  in  the  life  and  growth  of  the  tree  ;  and  these 
are  annually  renewed.     The  branches  of  each  year's  growth  are, 
therefore,  kept  in  fresh  communication,  by  means  of  the  newer 
layers  of  wood,  with  the  fresh  rootlets,  which  are  alone  active  in 
absorbing  the  crude  food  of  the  plant  from  the  soil.     The  fluid 
they  absorb  is  thus  conveyed  directly  to  the  branches  of  the  sea- 
son, which  develop  leaves  to  digest  it.     And  the  sap  they  receive, 
having  been  elaborated  and  converted  into  organic  nourishing 
matter,  is  partly  expended  in  the  upward  growth  of  new  branches, 
and  partly  in  the  formation  of  a  new  layer  of  wood,  reaching 
from  the  highest  leaves  to  the  remotest  rootlets. 

154.  Longevity  of  trees.     As  the  exogenous  tree,  therefore, 
annually  renews  its  buds  and  leaves,  its  wood,  bark,  and  roots, 
—  every  thing,  indeed,  that  is  concerned  in  its  life  and  growth,  — 
there  seems  to  be  no  necessary  cause,  inherent  in  the  tree  itself, 
why  it  may  not  h've  indefinitely.     Some  trees  are  known  to  have 
lived  for  one  and  two  thousand  years,  and  some  are  possibly 
older.1     Equally  long  may  survive  such  endogenous  trees  as  the 
Dragon  tree  (Dracaena) ,  which  have  provision  for  indefinite  in- 
crease in  diameter  (138),  and  for  the  production  of  branches. 
The  famous  Dragon  tree  of  Orotava,  in  Tenerifie,  now  destroyed 
by  hurricanes  and  other  accidents,  had  probably  reached  the  age 
of  more  than  two  thousand  years. 

155.  On  the  other  hand,  increase  in  height,  spread  of  branches 
and  length  of  root,  and  extension  of  the  surface  over  which  the 
annual  layer  is  spread,  are  attended  with  inevitable  disadvantage, 
which  must  in  time  terminate  the  existence  of  the  tree  in  a  way 
quite  analogous  to  the  death  of  aged  individual  animals,  which 
is  not  directly  from  old  age,  but  from  casualties  or  attacks  to 

1  The  subject  of  the  longevity  of  trees  has  been  discussed  by  DeCandolle, 
in  the  "  Bibliotheque  Universelle  "  of  Geneva,  for  May,  1831,  and  in  the  second 
volume  of  his  "  Physiologie  Vege'tale ; "  more  recently,  by  Alphonse  DeCan- 
dolle in  the  "  Bibliotheque  Universelle ; "  and  in  this  country  by  myself  in  the 
"  North  American  Review,"  for  July,  1844.  For  an  account  of  the  huge  Red- 
woods (Sequoias)  of  California,  see  Whitney's  Yosemite  Book. 


84  MORPHOLOGY  OF   STEMS. 

which  the  aged  are  either  increasingly  incident  or  less  able  to 
resist.  A  tree  like  the  Banyan  (59,  Fig.  71),  which  by  aerial 
roots  continues  to  form  new  trunks  for  the  support  and  sustenance 
of  the  spreading  branches,  and  thus  ever  advances  into  new  soil, 
has  a  truly  indefinite  existence ;  but,  then,  it  becomes  a  forest, 
or  is  to  be  likened  to  a  colony  propagated  and  indefinitely  in- 
creased by  suckers,  offsets,  or  other  subterranean  shoots.  So 
the  question  of  the  secular  continuation  of  the  individual  plant 
becomes  merged  in  that  of  continuation  of  the  race,  —  at  least  of 
a  bud-propagated  race,  —  the  answer  to  which  is  wholly  in  the 
domain  of  conjecture.1  However  this  may  be,  it  is  evident  that 
a  vegetable  of  the  higher  grade  is  not  justly  to  be  compared  with 
an  animal  of  higher  grade ;  that  individuality  is  incompletely 
realized  in  the  vegetable  kingdom ; 2  that  rather 

156.  The  Plant  is  a  Composite  Being,  or  Community,  lasting,  in 
the  case  of  a  tree,  through  an  indefinite  and  often  immense  num- 
ber of  generations.  These  are  successively  produced,  enjoy  a 
term  of  existence,  and  perish  in  their  turn.  Life  passes  onward 
continually  from  the  older  to  the  newer  parts,  and  death  follows, 
with  equal  step,  at  a  narrow  interval.  No  portion  of  the  tree  is 
now  living  that  was  alive  a  few  years  ago  ;  the  leaves  die  annu- 
ally and  are  cast  off,  while  the  internodes  or  joints  of  stem  that 
bore  them,  as  to  their  wood  at  least,  buried  deep  in  the  trunk 
under  the  wood  of  succeeding  generations,  are  converted  into 
lifeless  heart-wood,  or  perchance  decayed,  and  the  bark  that 
belonged  to  them  is  thrown  off  from  the  surface.  It  is  the  aggre- 
gate, the  blended  mass  alone,  that  long  survives.  Plants  of 
single  cells,  and  of  a  definite  form,  alone  exhibit  complete  indi- 
viduality; and  their  existence  is  extremely  brief.  The  more 
complex  vegetable  of  a  higher  grade  is  not  to  be  compared  with 
the  animal  of  the  highest  organization,  where  the  offspring  always 
separates  from  the  parent,  and  the  individual  is  simple  and  indi- 
visible. But  it  is  truly  similar  to  the  branching  or  arborescent 
coral,  or  to  other  compound  animals  of  the  lowest  grade,  where 
successive  generations,  though  capable  of  living  independently 
and  sometimes  separating  spontaneously,  yet  are  usually  devel- 
oped in  connection,  blended  in  a  general  body,  and  nourished 
more  or  less  in  common.  Thus,  the  coral  structure  is  built  up 
by  the  combined  labors  of  a  vast  number  of  individuals,  —  by 
the  successive  labors  of  many  generations.  The  surface  or  the 
recent  shoots  only  are  alive  ;  beneath  are  only  the  dead  remains 

1  See  Darwiniana,  xii.  338-356. 

2  As,  perhaps,  was  first  explicitly  stated  by  Engelmann,  in  his  inaugural 
essay,  De  Antholysi  Prodromus,  Introduction,  §  4. 


MORPHOLOGY  OF  LEAVES.  85 

of  ancestral  generations.  As  in  a  genealogical  tree,  only  the 
later  ramifications  are  among  the  living.  The  tree  differs  from 
the  coral  structure  in  that,  as  it  ordinarily  imbibes  its  nourish- 
ment mainly  from  the  soil  through  its  roots,  it  makes  a  downward 
growth  also,  and,  by  constant  renewal  of  fresh  tissues,  maintains 
the  communication  between  the  two  growing  extremities,  the 
buds  and  the  rootlets.  Otherwise,  the  analogy  of  the  two,  as  to 
individuality,  is  well-nigh  complete. 

SECTION  IV.     OF  LEAVES. 
§  1.   THEIR  NATURE  AND  OFFICE. 

157.  Leaf  (Lat.  Folium,  in  Greek  form  Phyllum),  as  a  botani- 
cal term,  has  on  the  one  hand  a  comprehensive,  on  the  other  a 
restricted  sense.     In  its  commonest  sense,  as  used  in  descriptive 
botany,  it  denotes  the  green  blade  only.     Yet  it  is  perfectly 
understood  that  the  footstalk  is  a  part  of  the  leaf,  and  therefore 
that  the  phrase  "  leaves  cordate,"  or  the  like,  is  a  short  way  of 
saying  that  the  blade  of  the  leaf  is  cordate  or  heart-shaped. 
Moreover,  two  appendages,  one  on  each  side  of  the  base  of  the 
footstalk,  when  there  is  any,  are  of  so  common  occurrence  that 
they  are  ranked  as  a  proper  part  of  the  organ.     So  that,  to  the 
botanist,  a  typical  leaf  consists  of  three  parts :    1 ,  BLADE  or 
LAMINA  ;    2,  FOOT-STALK  or  LEAF-STALK,  technically  PETIOLE  ; 
3,  A  pair  of  STIPULES.   (Fig.  142.) 

158.  The  blade,  being  the  most  important  part  of  an  ordinary 
leaf,  may  naturally  be  spoken  of  as  the  whole.     Petiole  and 
stipules  are  indeed  subsidiary  when  present,  and  are  not  rarely 
wanting.     Yet  sometimes  they  usurp  the  whole  function  of  foli- 
age, and  sometimes  there  is  no  such  distinction  of  parts. 

159.  Physiologically,  leaves  are  green  expansions  borne  by  the 
stem,  outspread  in  the  air  and  light,  in  which  assimilation  (3) 
and  the  processes  connected  with  it  are  carried  on.     Vegetable 
assimilation,  —  the  most  essential  function  of  plants,  being  the 
conversion  of  inorganic  into  organic  matter,  —  takes   place  in 
all  ordinary  vegetation  only  in  green  parts,  and  in  these  when 
exposed  to  the  light  of  the  sun.     And  foliage  is  an  adaptation 
for  largely  increasing  the  green  surface.     But  stems,  when  green, 
take  part  in  this  office  in  proportion  to  the  amount  of  surface, 
sometimes  monopolize  it,  and  in  various  cases  increase  their 
means  of  doing   so  by  assuming  leaf-like  forms.    (126-129.) 
Leaves,  especially  in  such  cases,  may  lose  this  function,  appear 
only  as  useless  vestiges,  or  may  be  subservient  to  various  wholly 


86  MORPHOLOGY  OP  LEAVES. 

different  uses.    Form  and  function,  therefore,  are  not  sure  indi- 
cations of  the  true  nature  of  organs. 

160.  Morphologically,  and  in  the  most  comprehensive  sense, 
leaves  are  special  lateral  outgrowths  from  the  stem,  definitely 
and  symmetrically  arranged  upon  it ;  in  ordinary  vegetation  and 
in  the  most  general  form  constituting  the  assimilating  apparatus 
(or  foliage) ,  but  also  occurring  in  other  forms  and  subserving 
various  uses.     Sometimes  these  uses  are  combined  with  or  sub- 
sidiary to  the  general  function  of  foliage  ;  sometimes  the  leaf  is 
adapted  to  special  uses  only.     So  the  botanist  —  recognizing  the 
essential  identity  of  organs,  whatever  their  form,  which  appear 
in  the  position  and  conform  to  the  arrangement  of  leaves  — 
discerns  the  leaf  in  the  cotyledons  of  a  bean  or  acorn,  the  scale  of 
a  lily-bulb  or  the  coat  of  an  onion,  the  scale  of  a  winter  bud,  and 
the  petal  of  a  blossom.     Therefore,  while  expanded  green  leaves 
(which  may  be  tautological^  termed  foliage-leaves)  are  taken  as 
the  proper  type,  the  common  name  of  leaves,  in  the  lack  of  any 
available  generic  word,  is  in  morphological  language  extended  to 
these  special  forms,  whenever  it  becomes  needful  to  express  their 
phylline  or  foliar  nature. 

161.  In  the  morphological  view,  all  the  plant's  organs  except- 
ing roots  (and  excepting  mere  superficial  productions,  such  as 
hairs,  prickles,  &c.),  belong  either  to  stem  or  to  leaves,  are 
either  cauline  or  phylline  in  nature.     To  the  latter  belong  all  the 
primary  outgrowths  from  nodes,  all  lateral  productions  which 
are  not  axillary.1    Whatever  is  produced  in  the  axil  of  a  leaf  is 
cauline,  and  when  developed  is  a  branch. 

162.  The  Duration  of  Leaves  is  transient,  compared  with  that 
of  the  stem.     They  may  be  fugacious,  when  they  fall  off  soon 
after  their  appearance ;  deciduous,  when  they  last  only  for  a 
single  season  ;  and  persistent,  when  they  remain  through  the  cold 
season,  or  other  interval  during  which  vegetation  is  interrupted, 
and  until  after  the  appearance  of  new  leaves,  so  that  the  stem  is 
never  leafless,  as  in  Evergreens.    In  many  evergreens,  the  leaves 
have  only  an  annual  duration  ;  the  old  leaves  falling  soon  after 
those  of  the  ensuing  season  are  expanded,  or,  if  they  remain 
longer,  ceasing  to  bear  any  active  part  in  the  economy  of  the 
vegetable,  and  soon  losing  their  vitality  altogether.     In  Pines 
and  Firs,  however,  although  there  is  an  annual  fall  of  leaves 
either  in  autumn  or  spring,  yet  these  were  the  produce  of  some 

1  There  are  cases  in  which  this  rule  is  of  difficult  application,  or  is  seem- 
ingly violated,  sometimes  by  the  suppression  of  the  subtending  leaf,  as  in  the 
inflorescence  of  Cruciferae,  rarely  in  other  ways,  to  be  explained  in  the 
proper  places. 


THEIR   STRUCTURE  AND   FORMS.  87 

season  earlier  than  the  last ;  and  the  branches  are  continually 
clothed  with  the  foliage  of  from  two  to  five,  or  even  ten  or  more 
successive  years.  On  the  other  hand,  it  is  seldom  that  all  the 
leaves  of  an  herb  endure  through  the  whole  growing  season,  the 
earlier  foliage  near  the  base  of  the  stem  perishing  while  fresh 
leaves  are  still  appearing  above.  In  our  deciduous  trees  and 
shrubs,  however,  the  leaves  of  the  season  are  mostly  developed 
within  a  short  period,  and  they  all  perish  in  autumn  nearly 
simultaneously. 

163.  Leaves  soon  complete  their  growth,  and  have  no  power 
of  further  increase.     Being  organs  for  transpiration,  a  very  large 
part  of  the  water  imbibed  by  the  roots  is  given  out  by  the  foliage, 
leaving  dissolved  earthy  matters  behind.     Assimilation  can  take 
place  only  in  fresh  and  vitally  active  tissue.     It  is  incident  to  all 
this  that  leaves  should  be  of  only  transient  duration,  at  least  in 
their  active  condition. 

164.  Defoliation.     The  leaves  of  most  Dicotyledons  and  some 
Monocotyledons  separate  from  the  stem  and  fall  by  means  of  an 
articulation  at  the  junction  with  the  stem,  which  begins  to  form 
early  in  the  season  and  is  completed  at  the  close.    There  is  a  kind 
of  disintegration  of  a  transverse  layer  of  cells,  which  cuts  off  the 
petiole  by  a  regular  line,  and  leaves  a  clean  scar,  such  as  is  seen 
in   Fig.    81,   85,   91.     Some   leaves,   notably  those  of  Palm?, 
Yucca,  and  other  endogens,  die  and  wither  on  the  stem,  or  wear 
away  without  falling. 

165.  In  temperate  climates,  defoliation  mostly  takes  place  at 
the  approach  of  winter.     In  warmer  climates  having  only  winter 
rain,  this  occurs  in  the  hot  and  dry  season. 

166.  Normal  Direction  or  Position.     The  leaf-blade  is  expanded 
horizontally,  that  is,  has  an  upper  and  an  under  surface.     When 
erect,  the  upper  surface  faces  the  axis  which  bears  it.     To  this, 
there  are  many  seeming  but  no  real  exceptions ;  that  is,  none 
which  are  not  explicable  as  deviations  or  changes  from  the  normal 
condition.   (213-217.) 

§  2.  THEIR  STRUCTURE  AND  FORMS  AS  ORGANS  OF  ASSIMILATION 
OR  VEGETATION,  i.  e.  AS  FOLIAGE. 

167.  The  Internal  Structure  or  Anatomy  of  the  leaf  needs  here 
to  be  examined  so  far  as  respects  its  obvious  parts  and  their 
general  composition.     The  leaf,  like  the  stem,  is  composed  of 
two  elements  (131),  the  cellular  and  the  woody.     The  cellular 
portion  is  the  green  pulp  or  parenchyma,  and  in  this  the  work 
of  assimilation  is  carried  on.     The  woody  is  the  fibrous  frame- 


88  MORPHOLOGY   OF   LEAVES. 

work,  the  separate  parts  or  ramifications  of  which  form  what  are 
variously  called  the  ribs,  —  a  sufficiently  proper  term,  —  nerves  or 
veins.  The  latter  names  may  suggest  false  analogies  ;  but  they 
are  of  the  commonest  use  in  descriptive  botany.  That  of  veins, 
and  of  its  diminutive,  veinlets,  for  the  smaller  ramifications,  is 
not  amiss ;  for  the  fibrous  framework  not  only  gives  firmness 
and  support  to  the  softer  cellular  apparatus,  i.  e.  forms  ribs, 
but  serves  in  the  leaf,  as  it  does  in  the  stem,  for  the  more  rapid 
conveyance  and  distribution  of  the  sap.  The  subdivisions  con- 
tinue beyond  the  limits  of  unassisted  vision,  until  the  fibro-vas- 
cular  bundles  are  reduced  to  attenuated  fibres  ramified  through 
the  parenchyma.  In  leafstalks,  the  woody  bundles  are  parallel, 
not  ramified,  and  arranged  in  varidus  ways  ;  in  Exogens  usually 
so  as  to  form  in  cross-section  an  arc  or  an  incomplete  or  com- 
plete ring.  In  leaves  serving  as  foliage  or  organs  of  assimilation, 
the  blade  is  the  important  part,  and  this  only  is  here  regarded. 

168.  The  characteristic  contents  of  these  cells  of  parenchyma 
are  grains  of  chlorophyll  (142s),  literally   leaf-green,  to  which 
the  green  color  of  foliage  is  wholly  owing,  and  which  may  be 
regarded   as  the  most  important   of  all   vegetable   products ; 
because  it  is  in  them  (or  in  this  green  matter,  whatever  its  form, 
189)  that  all  ordinary  assimilation  takes  place.     As  it  acts  only 
under  the  influence  of  light,  the  expanded   leaf-blade  may  be 
viewed  as  an  arrangement  for  exposing  the  largest  practicable 
amount  of  this  green  matter  —  the  essential  element  of  vegeta- 
tion —  to  the  light  and  air. 

169.  The  Parenchyma-cells,  constituting  the  green  pulp,  are 

themselves  arranged  in  accordance 
with  this  adaptation.  The  upper  stra- 
tum is  mostly  of  oblong  cells,  compactly 
arranged  in  one  or  more  layers,  their 
longer  diameter  perpendicular  to  the 
surface.  The  stratum  next  the  lower 
surface  of  the  leaf  consists  of  loosely 
arranged  cells,  with  longer  diameter 
usually  parallel  to  the  plane  of  the  leaf, 
often  irregular  in  form,  and  so  disposed 
as  to  leave  intervening  sinuous  air-spaces  freely  permeating  all 

FIG.  136.  A  magnified  section  through  the  thickness  of  a  leaf  of  Illicium  Flori- 
danum,  showing  the  irregular  spaces  or  passages  between  the  cells,  which  are  small  in 
the  upper  layer  of  the  green  pulp,  the  cells  of  which  (placed  vertically)  are  well  com- 
pacted, so  as  to  leave  only  minute  vacuities  at  their  rounded  ends;  but  the  spaces  are 
large  and  copious  in  the  rest  of  the  leaf,  where  the  cells  are  very  loosely  arranged : 
also  the  epidermis  or  skin  of  the  upper  (a)  and  of  the  lower  surface  of  the  leaf  (6), 
composed  of  perfectly  combined  and  thick- walled  empty  cells. 


THEIR   STRUCTURE  AND   FORMS.  89 

that  part  of  the  leaf.  (Fig.  136,  137.)  Hence  in  good  part  the 
deeper  green  hue  of  the  upper,  and  the  paler  of  the  lower  face 
of  leaves. 

170.  Epidermis.  The  whole  surface  of  leaves,  as  of  young 
stems,  is  invested  with  a  translucent  membrane,  composed  of 
one  or  sometimes  two  or  three  layers  of  empty  and  rather- 
thick-walled  cells.  This  is  the  skin  or  epidermis,  which  is  so 
readily  separable  from  the  succulent  tissue  of  such  leaves  as 
those  of  Stonecrop  and  other  species  of  Sedum.  It  is  of  a  single 
layer  in  the  Ilh'cium  (Fig.  136)  and  Lily  (Fig.  137)  ;  of  as 
many  as  three  in  the  firm  leaf  of  the  Oleander ;  is  generally 


137 

hard  and  thick  in  such  coriaceous  leaves  as  those  of  Pittosporum 
and  Laurustinus,  which  thereby  the  better  endure  the  dry  air  of 
rooms  in  winter. 

171.  Stomata  or  Breathing- pores.1  The  epidermis  forms  a 
continuous  protective  investment  of  the  leaf  except  where  certain 
organized  openings  occur,  the  stomata.  They  are  formed  by  a 
transformation  of  some  of  the  cells  of  the  epidermis  ;  and  consist 
usually  of  a  pair  of  cells  (called  guardian-cells) ,  with  an  opening 
between  them,  which  communicates  with  an  air-chamber  within, 
and  thence  with  the  irregular  intercellular  spaces  which  permeate 
the  interior  of  the  leaf.  Through  the  stomata,  when  open,  free 
interchange  may  take  place  between  the  external  air  and  that 

1  The  technical  name  lias  been  anglicized  stomates,  singular  stomate,  which 
has  no  advantage  over  the  proper  Greek,  sing,  stoma,  pi.  stomata. 

FIG.  137.  A  magnified  section  through  the  thickness  of  a  minute  piece  of  the  leaf 
of  the  White  Lily  of  the  gardens,  showing  also  a  portion  of  the  under  side  with  some 
breathing-pores,  stomata. 


90 


MORPHOLOGY   OP  LEAVES. 


within  the  leaf,  and  thus  transpiration  be  much  facilitated. 
When  closed,  this  interchange  will  be  interrupted  or  impeded. 
The  mechanism  of  stomata  is  somewhat 
recondite,  and  will  be  illustrated  in  the 
anatomical  and  physiological  volume  of 
this  series. 

172.  It  is  only  when  leaves  assume 

~V->  /P%    r\       a  vertical  or  edgewise  position  that  the 

(f  ~N  @p\J  \  stomata  are  in  equal  numbers  on  both 
faces  of  a  leaf.  Ordinarity,  the}r  occupy 
or  most  abound  on  the  lower  face,  which 
is  turned  away  from  the  sun  ;  but  in  certain  coniferous  trees  the 
reverse  of  this  is  true.  In  the  Water  Lilies  (Nymphaea,  Nuphar) , 
and  other  leaves  which  float 
upon  the  water,  the  stomata  all 
belong  to  the  upper  surface. 
Leaves  which  live  under  water, 
where  there  can  be  no  evapora- 
140  i«  tion,  are  destitute,  not  only  of 

stomata,  but  usually  of  a  distinct  epidermis  also.  The  number 
of  the  stomata  varies  from  800  to  about  170,000  on  the  square 
inch  of  surface  in  different  leaves.  In  the  Apple,  there  are  said 
to  be  about  24,000  to  the  square  inch  (which  is  under  the  average 
number,  as  given  in  a  table  of  36  species  by  Lindle}*)  ;  so  that 
each  leaf  of  that  tree  would  present  about  100,000  of  these 
orifices.  The  leaf  of  Dragon  Arum  is  said  to  have  8,000 
stomata  to  a  square  inch  of  the  upper  surface,  and  twice  that 
number  in  the  same  space  of  the  lower.  That  of  the  Coltsfoot 
has  12,000  stomata  to  a  square  inch  of  the  lower  epidermis, 
and  only  1,200  in  the  upper.  That  of  the  White  Lily  has 
from  20,000  to  60,000  to  the  square  inch  on  the  lower  sur- 
face, and  perhaps  3,000  on  the  upper;  and  they  are  so  re- 
markably large  that  they  may  be  discerned  by  a  simple  lens  of 
an  inch  focus. 

172.  Yenation,  the  veining  of  leaves,  &c.,  relates  to  the  mode 
in  which  the  woody  tissue,  in  the  form  of  ribs,  veins,  &c.,  is 
distributed  in  the  cellular.  There  are  two  principal  modes,  the 
parallel-veined  and  the  reticulated  or  netted-veined.  The  former 
is  especially  characteristic  of  plants  with  endogenous  stem  and 
monocotyledonous  embryo,  and  also  of  g3*mnospermous  trees, 

PIG.  138.  A  highly  magnified  piece  of  the  epidermis  of  the  Garden  Balsam,  with 
three  stomata  (after  Brongniart). 

FIG.  139.  Magnified  view  of  the  10,000th  part  of  a  square  inch  of  the  epidermis  of 
the  lower  surface  of  the  leaf  of  the  White  Lily,  with  Ita  stomata.  140.  A  single  stoma, 
more  magnified.  141.  Another  stoma,  widely  opea. 


THEIR   STRUCTURE  AND   FORMS. 


91 


which  have  exogenous  stems  and  at  least  dicotyledonous 
embryos.  The  latter  prevails  in  ordinary  plants  with  exogenous 
stem  and  dicotyledonous  embryo. 

173.  Parallel- veined  or  Nerved  leaves  (of  which  Fig.  143  is 
an  illustration)  have  a  framework  of  simple  ribs  (called  by  the 
earlier  botanists  nerves,  a  name  still  used  in  descriptions),  which 
run  from  the  base  to  tip,  or  sometimes  from  a  central  strong  rib 
to  margin  of  the  leaf,  in  a  generally  parallel  and  undivided  way, 
and  sending  off  or  connected  by  minute  veinlets  only.  Grasses, 


Lily  of  the  Valley,  and  the  like,  illustrate  the  commoner  mode 
in  which  the  threads  of  wood  run  from  base  to  apex.  The 
Banana  and  Canna  are  familiar  illustrations  of  a  mode  not  un- 
common in  tropical  or  subtropical  endogens,  in  which  the  threads 
or  "nerves"  run  from  a  central  rib  (midrib}  to  the  margin. 
Parallel- veined  leaves  are  generally  entire,  or  at  least  their 
margins  not  toothed  or  indented..  The  principal  exception  to 
this  occurs  when  the  ribs  or  the  stronger  ones  are  few  in  number 
and  radiately  divergent,  as  in  the  flabelliform  leaves  of  Fan- 
palms,  a  peculiar  modification  of  the  parallel- veined  type. 
Between  leaves  with  nerves  wholty  of  basal  origin,  and  those 
with  nerves  all  springing  from  a  midrib,  there  are  various  grada- 
tions, and  also  in  respect  to  curving.  But  parallel-veined  or 
nerved  leaves  may  be  classified  into 

FIG.  142.    A  leaf  of  the  Quince,  of  the  netted-veined  or  reticulated  sort,  with 
blade  (6),  petiole  or  leaf-stalk  (p),  and  stipules  {«<). 

FIG.  143.    Parallel-veined  leaf  of  the  Lily  of  the  Valley,  Convallaria  majalia. 


92  MORPHOLOGY  OF  LEAVES. 

Basal-nerved,  that  is,  with  the  nerves  all  springing  from  the 
base  of  the  leaf,  and 

Costal-nerved,  springing  from  a  midrib  or  costa.    Either  may  be 

Rectinerved,  the  nerves  running  straight  from  origin  to  apex 
or  margin  of  the  leaf,  as  the  case  may  be  ; 

Curvinerved,  when  curving  in  their  course,  as  in  the  leaves 
of  Funkia  and  in  Canna ; 

Flabellinerved,  where  straight  nerves  and  ribs  radiate  from  the 
apex  of  the  petiole,  as  in  Fan-palms  and  the  Gingko  tree. 

174.  In  typical  parallel- veined  leaves,  all  reticulation  is  con- 
fined to  minute  and  straight  cross- veinlets :  in  many,  these  are 
coarser,  branching,  and  reticulated ;  in  some,  as  in  Smilax  and 
Dioscorea,  only  the  primary  ribs  or  strongest  nerves  are  on  the 
parallel- veined  plan  ;  the  space  between  being  filled  with  reticu- 
lations of  various  strength  ;  thus  passing  by  gradations  into 

175.  Reticulated  or  Netted- veined  leaves.     In  most  of  these, 
from  one  to  several  primary  portions  of  the  framework  are 
particularly  robust,  and  give  origin  to  much  more  slender  ram- 
ifications, these  to  other  still  smaller  ones,  and  so  on.     The 
strong  primary  portions  are  RIBS  (costa)  ;  the  leading  ramifi- 
cations, VEINS  (venae)  ;  the  smaller  and  the  ultimate  subdivisions, 
VEINLETS  (venulce) .     All  or  some  of  the  veins  and  veinlets  are 
said  to  anastomose,  i.  e.  variously  to  connect  with  those  from 
other  trunks  or  ribs,  apparently  in  the  manner  of  the  veins  and 
arteries  of  animals,  forming  meshes.     But,  as  there  is  no  opening 
of  calibre  of  one  into  another,  the  word  is  etymologically  rather 
misleading.     More  properly,  it  is  said  that  the  veins  or  veinlets 
form  reticulations  or  net- work.     A  primary  division  of  reticulated 
leaves,  and  indeed  of  nerved  leaves  also,  into  two  classes,  is 
founded  upon  the  number  of  primary  ribs. 

176.  There  may  be  only  a  single  primary  rib  ;  this  traversing 
the  blade  from  base  to  tip  through  its  centre  or  axis  (as  in  Fig. 
142,  152-156)  is  called  the  MIDRIB.     There  may  be  others,  gen- 
erally few  (one,  two,  three,  or  rarely  four) ,  rising  from  the  apex 
of  the  petiole  on  each  side  of  the  midrib,  running  somewhat  par- 
allel with  it  or  more  or  less  diverging  from  it :  these  are  lateral 
ribs.     Among  parallel- veined  leaves,  the  Banana,  Canna,  &c., 
have  a  single  rib,  from  which  the  veins  (in  the  older  nomencla- 
ture here  called  nerves)  all  proceed.     Most  Lilies  and  the  like 
have  several  approximately  parallel  ribs,  but  the  midrib  pre- 
dominant: in  other  cases,  the  midrib  is  no  stronger  than  the 
others.     In  Fan-palms,  the  ribs  are  radiately  divergent,  giving 
a  fan-shaped  or  rounded  outline  to  the  blade.     In  reticulated 
leaves,  in  which  the  veins  all  spring  from  the  ribs,  tht  two 


THEIR    STRUCTURE   AND   FORMS. 


93 


classes  into  which  they  divide  are  the  pinnately  veined  and  the 
palmately  veined. 

177.  Pinnately   or    Feather- reined    (or    Penninerved)    leaves 
are  accordingly  those  of  which  the  veins  and  their  subdivisions 
are  side  branches  of  a  single  central  rib  (midrib),  which  traverses 
the  blade  from  base  to  apex ;  the  veins  thus  being  disposed  in 
the  manner  of  the  plume  on  the  shaft  of  a  feather.     (Fig.  142, 
152,  &c.)     Sometimes  these  continue  straight  and  undiminished 
from  midrib  to  margins  (straight- veined,  as  in  Beech  and  Chest- 
nut, Fig.  152),  sending  off  only  small  lateral  veinlets ;  some- 
times they  ramify  in  their  course  into  secondary  or  tertiary  veins, 
and  these  into  veinlets.     Pinnate  venation  in  reticulated  leaves 
naturally  belongs  to  leaves  which  are  decidedly  longer  than  wide. 

178.  Some  of  the  primary  veins,  commonly  among  the  lower, 
may  be  stronger  than  the  rest,  and  thus  take  on  the  character  of 
ribs,  or  by  gradations  pass  into  such.     The  leaf  of  the  common 
annual  Sunflower  (Fig.  155)  becomes  in  this  way  triple-ribbed  or 
tripli-nerved.     The  appearance  of  a  second  pair  of  such  strength 
ened  veins  makes  the  venation  quintupli-ribbed  or  quintupli-nerved. 


Through  the  approximation  of  such  strong  veins  to  the  base  of 
the  blade,  this  venation  may  pass  into  the 

179.  Palmately,  Digitately,  or  Radiately  Veined  (or  Palmi- 
nerved)  class,  of  which  leaves  of  common  Maples  and  the  Vine 
are  familiar  examples.  (Also  Fig.  158-160,  &c.)  In  these 

FIG.  144-157.  Various  forms  of  simple  leaves,  explained  in  the  text  and  in  the 
Glossary. 


94 


MORPHOLOGY   OP  LEAVES. 


there  are  three,  five,  seven,  or  sometimes  more  ribs  of  equal 
strength,  the  central  being  the  midrib,  and  each  with  its  system 
of  veins  which  ramify  and  form  meshes  in  the  interspaces.  Here 
the  whole  woody  portion  of  the  leaf  divides  equally  into  a  num- 
ber of  parts  upon  leaving  the  petiole  or  entering  the  blade.  The 
ribs  there  commonly  diverge  more  or  less  in  a  palmate  or  digitate 
manner  (i.  e.  like  outspread  fingers  of  the  hand,  or  the  claws  of  a 
bird,-  or  like  radii  of  more  or  less  of  a  circle)  :  so,  in  the  corre- 
lation of  outline  and  venation,  this  class  of  veining  goes  with 


roundish  circumscription.  This  is  not  so  true,  however,  in  a 
special  case,  viz.,  where  the  ribs,  however  divergent  below,  curve 
forward  and  all  run  to  the  apex  of  the  blade,  thus  imitating  the 
parallel-veined  system,  as  in  Rhexia  and  generally  in  the  family 
of  which  that  genus  is  the  single  northern  representative.1 

180.  Forms  as  to  Outline,  &c.  DeCandolle  conceived  the  shape 
of  leaves  (both  the  general  circumscription  and  the  special  con- 
figuration) to  depend  on  the  distribution  of  the  ribs  and  veins, 
and  quantity  of  the  parenchyma  in  which  these  were  outspread, 
— a  too  mechanical  view,  and  not  conformable  to  the  history  of 
development.  This  proves  that  the  framework  is  adapted  to  the 
parenchyma,  which  grows  and  shapes  the  organ  in  its  own  way, 
rather  than  the  parenchyma  to  it.  It  were  better  to  say  that  the 

1  In  Linnaean  terminology,  palmate  and  digitate  referred  to  particular  out- 
line only,  and  were  separately  used  to  denote  extent  of  division,— palmate,  not 
divided  down  to  the  petiole,  digitate,  when  divided,  like  the  claws  of  a  bird, 
quite  down  the  base.  DeCandolle  generalized  the  use  of  the  former  term, 
and  ever  since  the  two  have  been  used  interchangeably. 

fCO.  W8-166.    Various  forms  of  simple,  chiefly  palmately  veined  le»v««. 


THEIR   STRUCTURE   AND  FORMS. 


95 


two  elements  of  the  structure  are  correlated.  Descriptive  terms 
applied  to  leaves  are  equally  applicable  to  all  expanded  organs 
or  parts,  and  indeed  to  all  outlines.  Some  leading  forms  are 
here  enumerated  ;  and  all  are  defined  in  the  Glossary. 

181.  As  to  general  Circumscription,  proceeding  from  narrower 
to  broader  shapes,  and  then  to  those  with  either  narrowed  or 
notched  base,  leaf-blades  are 


Linear,  when  narrow,  several  times  longer  than  wide,  and  of 
about  the  same  breadth  throughout.  (Fig.  167.) 

Lanceolate,  or  Lance-shaped,  when  several  times  longer  than 
wide,  and  tapering  upwards  (Fig.  153,  168),  or  tapering  both 
upward  and  downward. 

Oblong,  when  nearly  twice  or  thrice  as  long  as  broad.  (Fig.  169.) 

Elliptical,  oblong  with  a  flowing  outline,  the  two  ends  alike 
in  width.  (Fig.  170.) 

Oval,  the  same  as  broadly  elliptical,  or  elliptical  with  the 
breadth  considerably  more  than  half  the  length. 

Ovate,  when  the  outline  is  like  a  section  of  a  hen's-egg  length- 
wise, the  broader  end  being  downward.  (Fig.  171,  155.) 

Orbicular,  or  Rotund,  circu- 
lar in  outline,  or  nearly  so. 
(Fig.  160.) 

Obovate,  inversely  ovate,  or 
ovate  with  the  narrower  end 
toward  the  base,  the  broader 
upward.  (Fig.  175,  145.) 

Cuneate,  or  Cuneiform,  that  is,  Wedge-shaped,  broad  above  and 
tapering  by  straight  lines  to  an  acute  base.  (Fig.  176,  148.) 

Spatulate,  rounded  above,  long  and  narrow  below,  like  a 
spatula.  (Fig.  174,  147.) 

Oblanceolate,  inverted  lance-shaped,  i.  e.  such  a  lanceolate 
leaf  as  that  of  Fig.  168,  but  with  the  more  tapering  end  at  base, 
as  in  Fig.  173.  To  those  who  restrict  the  term  lanceolate  to  the 
sense  of  a  narrow  leaf  tapering  equally  in  both  directions,  the 


FIG.  167-176.    Outlines  of  various  simple  leaves. 


MORPHOLOGY   OF   LEAVES. 


term  oblanceolate  is  superfluous.     The  following  terms  desig- 
nate leaves  with  a  notched  instead  of  narrowed  base. 

Cordate,  or  Heart-shaped,  when  a  leaf  of  an  ovate  form,  or 
something  like  it,  has  the  outline  of  its  rounded  base  turned  in 
(forming  a  notch  or  sinus)  where  the  stalk  is  attached,  as  in 
Fig.  172,  151.  Also  Fig.  149,  Pontederia,  a  leaf  of  the  parallel- 
veined  class. 

Reniform,  or  Kidney-shaped,  like  the  last,  only  rounder  and 
broader  than  long.  (Fig.  158.) 

Auriculate,  or  Eared,  having  a  pair  of  small  and  blunt  pro- 
jections, or  ears,  at  the  base,  as  in  Magnolia  Fraseri,  Fig.  178. 
Sagittate,  or  Arrow-shaped,  where  such  ears  are  acute  and  turned 

downwards,  while  the 
main  body  of  the  blade 
tapers  upwards  to  a 
point,  as  in  the  com- 
mon species  of  Sagit- 
taria  or  Arrow-head, 
and  in  the  Arrow- 
leaved  Polygonum. 
(Fig.  165,  177.) 

Hastate,  or  Halberd- 
shaped,  when  such 
lobes  at  the  base  point  outwards,  giving  the  leaf  the  shape  of  the 
halberd  of  the  olden  time,  as  in  Polygonum  arifolium  (Fig.  179) 
and  Sorrel,  Fig.  163. 

182.  Peltate  or  Shield-shaped  leaves  are  those  in  which  a  blade 
of  rounded  or  sometimes  of  other  shape  is  attached  to  the  petiole 
by  some  part  of  the  lower  surface,  instead  of  the  basal  margin : 
those  of  Water-shield  or  Brasenia,  of  Nelumbium,  and  of  Hydro- 
cotyle  umbellata  are  marked  examples.     The  anomaly  is  mor- 
phologically explained  by  a  comparison  with  deeply  cordate  or 
reniform  leaves  having  a  narrow  sinus,  such  as  those  of  Nym- 
phsea  or  Water  Lily,  and  by  supposing  a  union  of  the  approxi- 
mated edges  of  the  sinus.     Fig.  159  and  160,  from  two  species 
of  Hydrocotyle,  one  with  open  and  the  other  with  closed  sinus 
obliterated  by  the  union,  illustrate  this. 

183.  As  to  Extremity,  whether  base  or  apex,  there  are  several 
descriptive   terms,    expressive   of  the  principal  modifications ; 
such  as 

Acuminate,  tapering,  either  gradually  or  abruptly,  into  a 
narrow  more  or  less  prolonged  termination.  (Fig.  180.) 


FIG.  177-179.    Sagittate,  aurlculate,  and  hastate  leaves. 


THEIR   STRUCTURE   AND   FORMS. 


97 


Acute,  ending  in  an  acute  angle,  without  special  tapering,  as 
in  Fig.  181. 

Obtuse,  ending  with  a  blunt  or  roundish  extremity,  Fig.  182. 

Truncate,  with  termination  as  if  cut  off  by  a  straight  transverse 
line,  as  in  Fig.  183. 

Retuse,  with  an  obtuse  extremity  slightly  depressed  or  re-enter- 
ing, as  in  Fig.  184. 

Emarginate,  with  a  more  decided  terminal  notch,  Fig.  185. 

Obcordate,  inversely  heart-shaped,  i.  e.  like  cordate,  but  the 
broader  end  and  its  strong  notch  at  apex  instead  of  base,  Fig. 
186.  This  and  the  following  terms  are  applicable  to  apex  only. 

Mucronate,  abruptly  tipped  with  a  small  and  short  point,  like  a 
projection  of  the  midrib,  as  in  Fig.  188. 

Cuspidate,  tipped  with  a  sharp  and  rigid  point,  as  in  Fig.  187. 


184.  As  to  Margin  or  special  Outline,  the  terminology  proceeds 
upon  the  convenient  supposition  of  a  blade  with  quite  entire  margin, 
but  subject  to  incisions,  which  give  rise  to  notches  or  clefts,  if  we 
regard  the  sinuses  ;  or  to  teeth,  lobes,  segments,  &c.,  if  we  regard 
the  salient  portions  between  the  sinuses.    The  ribs,  or  the  stronger 
veins,  &c.,  commonly  terminate  in  the  teeth  or  lobes  ;  but  in  Cicuta 
maculata,  and  in  a  few  other  cases,  they  run  to  the  notches. 

185.  Dentation  relates  to  mere  marginal  incision,  not  extend- 
ing deeply  into  the  blade.     The  blade  is  said  to  be 

Entire,1  when 
the  margin  is  com- 
pletely filled  out 
to  an  even  line,  as 
in  Fig.  173-179. 

Serrate,  when 
with  small  and 
sharp  teeth  direct-  j 
ed  forward,  like 
the  teeth  of  a  saw,' 
as  in  Fig.  189. 

Serrulate  is  the     m         190         m         192  193  m 

diminutive  of  serrate,  and  is  equivalent  to  minutely  serrate. 

1  Integerrimus-a-um,  or  quite  entire,  is  the  term  in  Latin  terminology.     Inte 
ger  means  undivided  or  not  lobed. 

FIG.  180-188.    Terminations  of  leaves. 

FIG.  189-191    Dentation  of  pinnately  veined  leaves. 

7 


98  MORPHOLOGY   OP  LEAVES. 

Dentate,  or  Toothed,  a  general  term  for  toothing,  specially 
applied  to  the  case  of  salient  teeth  which  are  not  directed  for- 
ward or  towards  the  apex  of  the  blade,  Fig.  190. 

Crenate,  or  Scalloped,  the  same  as  dentate  or  serrate,  but  with 
teeth  much  rounded,  Fig.  191. 

Repand,  or  Undulate,  when  the  margin  is  a  wavy  line,  bending 
slightly  inward  and  outward,  Fig.  192. 

Sinuate,  when  this  wavy  line  is  stronger  or  distinctly  sinuous, 
as  in  Fig.  193. 

Incised,  when  cut  by  sharp  and  irregular  incisions  more  or  less 
deeply,  Fig.  194.  This  is  intermediate  between  dentation  and 

186.  Lobation  or    Segmentation.     When   the   blade  is  more 
deeply  penetrated  by  incisions  from  the  margin,  that  is,  when 
the  spaces  between  the  ribs  or  principal  veins  are  not  filled  to 
near  the  general  outline,  it  is  said  to  be  lobed,  cleft,  parted,  or 
divided,  according  to  the  degree  of  separation  ;  and  the  portions 
are  called  lobes,  segments,  divisions,  &c.     The  most  general  name 
for  such  parts  of  any  simple  blade  is  that  of  lobes.     More  par- 
ticularly a  leaf-blade,  or  other  body,  is  said  to  be 

Lobed,  when  the  division  extends  not  more  than  half  way  down, 
and  either  the  sinuses  or  the  lobes  are  rounded  ; 

Cleft,  when  the  division  is  half  way  down  or  more,  and  the 
lobes  or  sinuses  narrow  or  acute  ; 

Parted,  when  the  divisions  reach  almost,  but  not  quite,  to  the 
base  or  the  midrib ; 

Divided,  when  they  sever  the  blade  into  distinct  parts,  which 
makes  the  leaf  compound.  (193.) 

187.  LOBE  is  the  common  name  of  one  of  the  parts  of  a  simple 
blade,  especially  when  there  is  only  one  order  of  incision.     But 
when  there  are  more,  as  when  a  leaf  is  divided  or  parted  and 
these  primary  lobes  again  lobed  or  cleft,  the  lobes  of  first  order 
are  commonly  called  SEGMENTS  (sometimes  divisions  or  partitions), 
and  the  parts  of  these,  Lobes.     Or  the  lobes  may  be  designated 
as  primary,  secondary,  tertiary,  &c.     Ultimate  portions  or  small 
lobes  may  be  called  Lobules  or  Lobelets.     Also  the  portions  of  a 
quite  divided  blade  take  the  name  of  Lea/lets.     By  proper  selec- 
tion of  terms,  the  degree  of  division  or  lobing  may  thus  be 
expressed  in  a  single  word. 

188.  As  to  Number  of  parts,  this  may  be  tersely  expressed  by 
combination  with  the  adjective  term  applicable  to  the  degree  ;  as, 
Two-lobed,  Three-lobed,  Five-lobed,  Many-lobed,  &c.  ;  or  Two-Five- 
cleft,  Many-cleft,  &c.,  in  Latin  form  Bifid,  Trifid,  Muhifid,  &c.  ; 
Two-five-parted,    &c.,    according  to   the   number  of  divisions 
which  extend  almost  to  the  base  or  axis  ;  Two-Five-divided  (in 


THEIR  STRUCTURE  AND  FORMS. 


99 


Latin  form  Bisected,  Trisected,  &c.),  when  there  are  two  or  three 
or  more  complete  divisions  of  the  blade. 

189.  As  to  Arrangement  of  parts,  this  may  be  simply  and  best 
expressed  by  taking  into  account  the  nature  of  the  venation  or 
the  distribution  of  the  ribs,  &c.,  which  controls  or  is  co-ordinated 
with  the  disposition  of  the  lobes.  Pinnately  veined  leaves, 
when  lobed,  must  needs  have  the  incisions  directed  to  the  mid- 
rib ;  palmately- veined  or  radiated,  to  the  apex  of  the  petiole. 
*The  lobes  or  divisions  of  the  first  will  be  pinnately,  of  the  second 


195  196 


palmately  disposed.  Accordingly,  the  three  leaves  of  as  many 
species  of  Oak,  Fig.  195,  196,  and  197,  represent  respectively  a 
pinnately  lobed,  pinnately  cleft,  and  pinnately  parted  leaf,  while  the 
accompanying  leaf  of  Celandine,  Fig.  198,  is  pinnately  divided. 
The  first  three,  however,  when  the  degree  of  incision  is  not  par- 
ticularly in  question,  usually  pass  under  the  common  term  of 
pinnatifid,  Fig.  195  moderately,  Fig.  197  deeply.  The  number 
of  lobes,  when  definitely  marked,  may  come  into  the  descriptive 
phrase,  as  pinnately  1-lobed,  pinnately  1-cleft,  parted,  or  divided, 
as  the  case  may  be. 

190.  Similarly,  Figures  199  to  202  represent,  respectively,  a 
palmately  three-lobed,  three-cleft,  three-parted,  and  three-divided,  or, 
in  Latin  form,  trilobate,  trifid,  tripartite,  and  trisect  or  trisected 
leaf.  Fig.  166  is  a.  palmately  ^-parted  leaf;  Fig.  164,  palmately 


FIG.  195-198.    Pinnately  lobed,  cleft,  parted,  and  divided  leaves. 
FIG.  199-202.    Palmately  3-lobed,  cleft,  parted,  and  divided  leave 


100  MORPHOLOGY  OF  LEAVES. 

muUifid,  &c.  Fig.  162,  a  leaf  of  Dragon  Arum,  is  palmately 
^-parted.  But,  as  the  lateral  sinuses  are  not  so  deep  as  the 
others,  the  leaf  is  said  to  be  pedately  parted,  or  pedate,  in  the 
early  terminology. 

191.  Moreover,  as  the  lobes  or  divisions  of  a  leaf  may  be 
again  similarly  lobed  or  parted,   &c.,  this  composition  may  be 
indicated  by  the  prefix  twice,  thrice,  &c.,  as  twice  pinnatifid  or 
Upinnatijid,  thrice  pinnately  parted,   thrice  palmately  parted,  and 
the  like.     Thus,  a  word  or  two,  or  a  short  phrase,  may  describe 
even  a  complex  leaf,  so  as  to  convey  a  perfectly  clear  and  defi- 
nite idea  of  its  conformation. 

192.  A  distinction  should  now  be  drawn  between  simple  and 
compound  leaves.     The  distinction  cannot  be  both  natural  and 
absolute  ;  for  the  one  may  pass  variously  into  the  other.     Simple 
leaves,  which  have  been  thus  far  considered,  have  a  single  lamina 
or  blade,  which  may,  however,  at  one  extreme  be  entire,  at  the 
other  many-parted,  and  even  several  times  divided. 

193.  Compound  Leaves   are  those   which  have  from  two  to 
many  distinct  blades,  on  a  common  leafstalk.     These  blades, 
called  LEAFLETS,  may  be  sessile  on  the  common  leafstalk,  or  they 
may  have  leafstalks  of  their  own.     As  the  leaf  very  commonly 
separates  in  age  by  an  articulation  of  its  petiole  with  the  stem, 
so  leaflets  are  commonly  more  or  less  articulated  with  the  com- 
mon petiole.     When  the  leaf,  with  its  petiole,  falls  from  the 
stem,  the  leaflets  may  as  completely  separate  from  the  common 
petiole.     They  do  not  always  do  this.     Divided  leaves,  such  as 
those  of  Fig.   198  and  202,  though  ranked  among  the  simple 
sorts,   are  compound   in  the  sense  of  having  distinct  blades, 
but  without  articulation.     Some  of  these  blades  are  apt  to  be 
confluent ;  that  is,  a  divided  leaf  is  often  in  part  merely  parted, 
as  in  the  upper  portion  of  Fig.  198.     Such  leaves  are  so  inter- 
mediate between  simple  and  compound  that  it  becomes  indiffer- 
ent, or  a  matter  of  convenience  to  be  settled  by  analogy,  under 
which  head  or  by  what  language  they  shall  be  described.     How- 
ever, most  leaves  are  so  constituted  as  to  leave  no  doubt  whether 
they  are  simple  or  compound. 

194.  The  leaflets  of  a  compound  leaf  being  homologous  with 
the  lobes  or  segments  of  a  simple  leaf,  indeed  being  such  segments 
fully  isolated,  the  two  sorts  fall  under  the  same  types.     A  pin- 
nately veined  simple  leaf  is  the  homologue  of  one  kind  of  com- 
pound leaf;    a   radiately  veined   leaf,  of  the  other.      That  is, 
compound  leaves  are  either  pinnate  or  palmate. 

195.  Pinnate  Leaves  (Fig.  203-205)  are  those  in  which  the 
leaflets  are  arranged  along  the  sides  of  a  petiole,  or  rather  of  its 


THEIR   STRUCTURE   AND   FORMS. 


101 


prolongation,  the  RHACHIS,  which  answers  to  the  midrib  of  a  pin- 
nately  veined  simple  leaf.  There  are  three  principal  sorts,  and 
some  subordinate  ones.  That  is,  a  pinnate  leaf  may  be 


Impari-pinnate,  or  pinnate  with  an  odd  leaflet,  i.  e.  a  terminal 
one,  as  in  Fig.  203  ;  and  this  is  the  commoner  case. 

Cirrhiferous  Pinnate,  or  pinnate  with  a  tendril  (Fig.  204),  as 
in  the  proper  Pea  tribe  and  Bignonia.  Here  either  the  termi- 
nal leaflet  only,  or  the  upper  lateral  leaflets  also,  are  replaced 
by  tendrils. 

Pari-pinnate,  or  Abruptly  Pinnate,  destitute  of  a  terminal  leaflet 
or  of  any  thing  answering  to  it,  as  in  Fig.  205. 

Interruptedly  Pinnate  denotes  merely  a  striking  inequality  of 
size  among  the  leaflets  :  Lyrately  Pinnate,  one  in  which  the  termi- 
nal leaflet  is  largest  and  the  lower  small. 

196.  Palmate  or  Digitate  Leaves  (Fig.  206,  93)  are  those  in 
which  the  leaflets  all  stand  on  the 
summit  of  the  petiole.  Digitate 
(fingered)  was  the  old  name,  when 
the  term  palmate  was  restricted  to 
a  simple  but  palmately  lobed  leaf 
of  this  type.  But  since  the  time 
of  DeCandolle  the  two  names  have 
been  used  interchangeably.  Pal- 
mate leaves  have  no  primary  dis- 
tinction into  sorts,  except  as  to 
the  number  of  leaflets.  These  can 
never  be  very  numerous ;  but  there  are  fully  a  dozen  in  some 

FIG.  203.  An  impari-pinnate  or  odd  pinnate  leaf.  204.  Pinnate  with  a  tendril 
205.  Abruptly  pinnate  leaf  of  a  Cassia. 

FIG.  206.    Palmately  or  digitately  5-foliolat«  leaf  of  a  Buckeye,  ^Esculus. 

•TAV 
SANTA  BAnb/>.aA.  CALIFORNIA 


102  MORPHOLOGY  OF  LEAVES. 

Lupines.     More  commonly  there  are  only  five  to  nine,  or  only 
three,  rarely  two,  or  even  a  single  one. 

197.  Number  of  leaflets  may  be  indicated  by  an  adjective 
expression  composed  of  the  proper  Latin  numeral  prefixed  to 
foliolate  (Foliolum,  diminutive  of  folium,  answering  to  leaflet) . 
Thus,   bifoliolate,  of  two  leaflets;  trifoliolate,  of  three  leaflets; 
quadrifoliolate,  of  four  ;  quinquefoliolate,  of  five  :  plurifoliolate,  or 
multifoliolate,  of  several  or  numerous  leaflets,  &c.     These  terms 
are  still  more  descriptive  when  accompanied  by  the  word  pin- 
nately  or  palmately,  indicative  of  the  kind  of  compound  leaf;  as, 
palmately  or  digitately  trifoliolate  (common  Clover-leaf,  Fig.  211), 
or  5-foliolate,  as  in  Buckeye  (Fig.  206),  and  so  on.    Also,  pinnately 
IG-foliolate,   as   in   Fig.    205,  or    17-foliolate,   as  in  Fig.  203 ; 
pinnately  trifoliolate,  as  in  Phaseolus,  and  in  the  low  Hop-Clover, 
Trifolium  procumbens.1 

198.  But,  in  either  class  of  compound  leaves,  the  leaflets  may 
be  reduced  to  a  minimum  number.     A  pinnately  trifoliolate  leaf 
is  one  of  the  impari-pinnate  kind  reduced  to  three  leaflets,  to  one 
pair  and  the  odd  one  ;  and  this  is  distinguished  from  a  palmately 
trifoliolate  leaf  by  the  attachment  of  the  pair  at  some  distance 
below  the  apex  of  the  petiole,  and  by  the  articulation  above  this, 
which  marks  the  junction  of  the  terminal  leaflet's  petiole  (or  its 
base,  if  sessile)  with  the  rhachis  or  common  petiole. 

199.  Unifoliolate  compound  leaves  (by  no  means  a  direct  con- 

tradiction in  terms)  are  by  this  articulation  distin- 
guished from  simple  leaves  which  they  simulate. 
See  the  leaf  of  the  common  Barberry,  Fig.  207. 
In  other  species,  of  the  Mahonia  section,  the  leaves 
are  all  pinnately  3-9 -foliolate,  with  well-developed 
common  petiole :  in  the  true  Berberis,  they  are  all 
thus  reduced  to  the  terminal  and  long-petiolulate 
leaflet,  on  an  almost  obsolete  petiole.  Orange  and 
Lemon  leaves  are  in  similar  case,  but  with  the  joint 
close  to  the  blade.  A  comparison  with  near  rela- 
tives shows  that  these  are  also  unifoliolate  leaves 

of  the  pinnate  kind ;  though  this  could  not  be  ascertained  by 

inspection. 

200.  Decompound  or  Twice  and  Thrice  Compound  Leaves.  These 
are  to  once  pinnate  or  once  palmate  leaves  what  the  latter  are  to 

1  In  pinnate  leaves,  each  leaflet  usually  has  its  opposite  fellow,  and  the 
number  may  be  \ndicated  by  the  pairs,  as  unijugate,  bijugate,  trijugate,  and 
pluri jugate,  according  to  the  number  ofjuga,  or  pairs. 

FIG.  207.  Unifoliolate  leaf  of  Berberis  vulgaris,  with  partial  petiole  articulated  to 
the  extremely  aha  t  true  petiole. 


THEIR   STRUCTURE  AND   FORMS.  103 

simple  leaves.  As  leaflets  may  be  toothed,  lobed,  or  parted,  so 
what  answers  to  a  single  leaflet  may  appear  as  leaflets  of  a  second, 
or  again  of  a  third,  or  even  of  a  fourth  order.  Decompound  is  a 
good  general  name  for  all  more  than  once  compounded  leaves ; 


but  the  name  has  been  applied  rather  to  irregularly  many-times 
parted  or  dissected  leaves  (such  as  those  of  Dicentra) ,  or  to 
those  more  than  thrice  compounded.  Of  regularly  twice  or 
thrice  compound  leaves,  the  commonest  are  the 

Bipinnate  or   Twice   Pinnate,  of  ordinary 
occurrence  in  the  Mimoseous  and  Csesalpi- 
neous,  but  not  in  the  Papilionaceous,  Legu- 
minosse.    Fig.  208  represents  a  bipinnate  leaf 
of  the  Honey  Locust  (Gleditschia) ,  with  the 
variation  (common  with  that  tree)  that  some 
of  the  partial  petioles,  in   this  figure  only  the 
lowest,  bears  a  single  leaflet,  while  the  others 
are  extended  into   secondary  rhachises 
nished  with  numer- 
ous leaflets,  mostly 
in  the  abruptly  pin- 
nate style.     On  the 
same  tree,  the  earlier 
leaves,    which    are 
clustered    on   short 
spurs,     are    simply 
pinnate.    The  large 
leaves  of  Gymnocladus  are  similarly  and  abruptly  bipinnate, 

FIG.  208.    A  bipinnate  and  multifoliolate  leaf  of  Gleditschia  or  Honey  Locust. 
FIG.  209.    Bipinnate  leaves  of  Sensitive  Plant,  Mimosa  pudlca,  with  approximate 


104  MORPHOLOGY   OF    LEAVES. 

except  at  the  base,  which  is  simply  pinnate  or  with  one  or  two 

pairs  of  simple  leaflets. 

Tripinnate  or  Thrice  Pinnate  leaves  of  a  regular  sort  are  rare ; 

but,  with  some  irregularity,  they  occur  in  many  species,  as  in 

Aralia,  &c.     This  extent  of  division,  and  even  much  greater,  is 

common  in  Ferns. 
Digitate-Pinnate  is  where  the  primary  division  of  the  petiole  is 

on  the  palmate  or  digitate  plan  ;  the  secondary,  on  the  pinnate. 

This  seems  to  be  the  case  in  the  Sensitive  Plant,  Mimosa  pudica, 

Fig.  209.     But  the  leaf  is  here  truly  bipinnate  with  the  primary 

divisions  very  crowded  at  the  apex  of  petiole. 

Conjugate- Pinnate  is  the  same  arrangement,  with  the  primary 

divisions  a  single  pair,  at  the  apex  of  the  petiole,  and  the  leaflets 

pinnately  arranged  on  these. 

Digitately  or  Palmately  Decompound  in  a  nearly  regular  way 
is  not  an  uncommon  case. 
Usually,  the  petiole  is  succes- 
sively three-forked,  as  in  Fig. 
210,  when  the  leaf  is  said  to 
be  biternate  (twice  ternate), 
triternate  (thrice  ternate),  or 
quadriternate  (four  times  ter- 
nate), &c.,  according  to  the 
number  of  times  it  divides,  or 
2— 3— 4^-times  ternately  compound. 
no  ^  The  ultimate  divisions  in  such 

cases  of  threes  are  commonly  of  the  pinnately  trifoliolate  type. 

201.  Pinnae  is  a  convenient  name  for  the  partial  petioles  of  a 
bipinnate  leaf,  taken  together  with  the  leaflets  that  belong  to 
them.     Thus,  the  Sensitive  Plant,  Fig.  209,  has  four  pinnae,  or 
two   pairs ;   the   Honey  Locust,   Fig.  208,  a   greater  number. 
When  such  leaves  are  still  further  compounded,  the  pinnae  of 
higher  order,  or  the  ultimate  ones,  take  the  diminutive  term  of 
PINNULE  or  PINNULES.     The  blades  these  bear  are  the  Leaflets. 

202.  The  Petiole  or  Leafstalk  is  a  comparatively  unessential 
part  of  the  leaf.     It  is  often  wanting  (then  the  blade  is  sessile)  ; 
it  may  be  absent  even  in  compound  leaves  of  the  palmate  type, 
the  leaflets  rising  side  by  side  from  the  stem.     When  present,  it 
is  usually  either  round,  or  half-cylindrical  and  channelled  on  the 
upper  side.     In  the  Aspen,  it  is  flattened  at  right  angles  with 
the  blade,  so  that  the  slightest  breath  of  air  puts  the  leaves  in 
motion.     Sometimes  it  is  much  dilated  and  membranaceous  at 

FIG.  210.    Quadri-ternately  compound  or  ternately  decompound  leaf  of  Thalictrum 
Cornuti. 


THEIR   STRUCTURE   AND   FORMS.  105 

base,  as  in  many  umbelliferous  plants ;  sometimes  it  forms  a 
sheath,  occasionally  it  is  bordered  with  appendages,  &c.  Peti- 
oles may  assume  special  functions,  to  be  hereafter  considered. 
The  woody  and  vascular  tissue  runs  lengthwise  through  the 
petiole,  in  the  form  usually  of  a  definite  number  of  parallel 
threads,  to  be  ramified  in  the  blade.  The  ends  of  these  threads 
are  apparent  on  the  base  of  the  leafstalk  when  it  falls  off,  and  on 
the  scar  left  on  the  stem,  as  so  many  round  dots  (Fig.  81,  85, 
91),  of  a  uniform  number  and  arrangement  in  each  species. 

203.  Partial  Petioles  are  the  divisions  of  the   petiole   in   a 
compound  leaf.     The  footstalk  of  a  leaflet  takes  the  diminutive 
name  of  PETIOLULE. 

204.  Stipules  (157)  are  lateral  appendages,  one  each  side  of 
the  base  of  the  petiole,  sometimes  free 

from  it  and  from  each  other  (Fig. 
142),  sometimes  attached  by  one 
edge  to  its  base  (Fig.  211),  some- 
times united  with  each  other  into  a 
single  body  (Fig.  212)  in  various 
ways  or  degrees.  In  the  latter  case, 
they  usually  appear  to  be  within  the 
base  of  the  leaf  or  leafstalk ;  or,  as 
in  the  Plane-tree,  they  may  be  joined 
into  one  over  against  the  leaf,  as 
if  opposite  to  it,  but  their  normal 
position  is  supposed  to  be  lateral  or 
marginal  to  the  petiole.  Sometimes 
they  are  foliaceous  in  appearance  and 
in  function ;  sometimes  they  are  dry 
and  colorless  or  scale-like,  reduced  to 
mere  epidermal  tissue,  and  evidently 
functionless  ;  sometimes  (as  in  Mag- 
nolia, Fig.  81,  Fig-tree,  and  Beech), 
they  serve  as  bud-scales,  and  fall  when 
the  leaves  develop  ;  sometimes  they  are 
reduced  to  a  mere  bristle,  or  take  the 
form  of  a  spine,  as  in  the  Locust  (Robinia) .  Between  salient 
expansions  or  wing-like  margins  of  the  base  of  the  petiole, 
such  as  those  of  the  Saxifrage  tribe,  and  stipules  adnate  to  the 
margins  of  the  petiole,  as  in  most  Rosacese,  there  is  no  clear 
limitation.  But  presence  or  absence  of  stipules  generally  runs 

FIG.  211.  Clover-leaf,  with  adnate  stipules.  212.  Ochreate  stipules  (ochrea  or  ocreal 
of  Polygoaum  orientale,  sheathing  the  stem  for  some  distance,  and  ending  in  a  spread- 
ing border. 


106  MORPHOLOGY   OP  LEAVES. 

through  a  natural  order.  Yet  what  are  called  stipules  in  one  order 
may  pass  for  expansions  or  appendages  of  the  petiole  in  another. 
In  Spergularia,  some  stipules  are  connate  around  the  base  of  the 
pair  of  leaves,  including  them  as  well  as  the  stem  in  the  sheath.1 

205.  Stipules,  which  are  normally  a  pair,  may  unite  into  one 
body,  either  adnate  to  the  inner  face  of  the  leaf,  as  in  some 
species  of  Potamogeton,  or  united  opposite  the  leaf,  as  in  Plane- 
tree,  or  united  inter  se  in  a  sheath,  as  in  Polygonum.    Also  when 
the  leaves  are  opposite  and  the  stipules  thus  brought  into  prox- 
imity, the  adjacent  half  stipules  of  the  two  leaves  may  coalesce, 
and  present  the  appearance  of  only  two  stipules  to  two  leaves, 
as  in  many  Rubiaceae.     A  notch  or  fork   at  the  apex  often 
indicates  the  composition. 

206.  Sheathing  stipules,  like  those  of  Polygonum  (Fig.  212), 
are  said  to  be  ochreate,  or   (better)   ocreate ;  the  sheath,  thus 
likened  to  a  leggin  or  the  leg  of  a  boot,  is  an  OCHREA,  as  written 
by  Willdenow,  or  better  OCREA. 

207.  The  LIGULE  of  Grasses  (Fig.  150)  is  seemingly  a  thin  and 
scarious  extension  of  the  lining  to  the  sheath  which  answers  to 
petiole  in  such  leaves :  it  projects  at  the  junction  of  the  sheath 
and  blade,  there  forming  a  kind  of  ocrea ;  and  it  is  generally 
regarded  as  a  sort  of  stipule. 

208.  Stipels  (Stipetta)  are  as  it  were  stipules  of  leaflets,  which 
are  common  in  certain  tribes  of  Papilionaceous  Leguminosse,  e.  g. 
in  the  Phaseoleae,  in  Wistaria,  Locust,  &c. ;  also  in  Staphylea. 
They  are  small  and  slender,  and,  unlike  stipules,  they  are  single 
to  each  leaflet,  except  to  the  terminal  one,  which  has  a  pair.   As 
leaves  furnished  with  stipules  are  said  to  be  stipulate,  so  leaflets 
with  stipels  are  stipellate. 

209.  Some  unusual  modifications  of  leaves  as   foliage.       In 
leaves  as  illustrated  thus  far,  it  is  the  lamina  or  blade  which  is 
expanded  to  do  the  work  of  foliage ;  which  is  expanded  hori- 
zontally, so  as  to  present  upper  and  under  surfaces,  one  to  the 
sky,  the  other  to  the  ground ;  which  is  bilaterally  symmetrical 
or  substantially  so,  the  two  lateral  halves  being  nearly  if  not 
quite  alike  ;  and  which  is  affixed  to  the  stem  at  the  basal  margin, 
or  some  part  of  it,  with  or  without  a  petiole.     Various  deviations 
or  apparent  deviations  from  this  pattern  occur.    Some  of  them  are 
of  comparatively  small  account  and  simple  explanation,  such  as 

210.  Inaeqnilateral  Leaves,  being  unsymmetrical  by  the  much 
greater  development  of  one  side.     This  is  illustrated   in   the 
whole  genus  Begonia  (as  in  Fig.  161),  consisting  of  many  spe- 

1  As  pointed  out  by  Prof.  A.  Dickson,  in  Nature,  xviii.  607. 


THEIR  STRUCTURE  AND  FORMS. 


107 


cies,  some  of  which  are  moderately,  and  most  of  them  strikingly, 
oblique  in  this  way.  Elm-leaves,  and  the  like,  are  more  or  less 
inequilateral  at  the  base. 

211.  Connate  and  Perfoliate  Leaves.  These  are  explained  by 
the  union  of  con- 
tiguous leaf-edges. 
Peltate  leaves,  to 
which  a  paragraph 
has  already  been 
given  (182),  come 
under  the  same 
head ;  the  seeming 
attachment  of  the 
petiole  to  the  lower 
face  of  the  blade 
being  the  result  of 
a  congenital  union 
of  the  edges  of  the 
sinus.  In  a  sessile 
leaf,  when  such  a  213 

union  takes  place,  it  surrounds  and  encloses  that  portion  of  the 
stem  ;  which  is  thus  perfoliate.  (Fig. 
213,  214.)  It  is  the  stem  which  is 
literally  perfoliate,  i.e.  which  seem- 
ingly passes  through  the  leaf ;  but  it 
is  customary,  though  etymologically 
absurd,  to  call  this  a  perfoliate  leaf! 
Uvularia  perfoliata  (Fig.  213),  in  the 
later  growth  of  the  season,  reveals  the 
explanation  of  the  perfoliation :  the 
base  of  the  lower  leaves  conspicuously 
surrounds  and  encloses  the  stem  :  that 
of  the  upper  is  merely  cordate  and 
clasping ;  the  uppermost  simply  ses- 
sile by  a  rounded  base.  Baptisia 
perfoliata  (Fig.  214)  is  a  more 
strongly  marked  case  of  perfoliation. 
But  there  are  good  morphological 
reasons  for  inferring  that  this  seemingly  simple  leaf  consists  of 
a  pair  of  stipules  and  a  leaflet  combined.  An  occasional  mon- 
strosity verifies  this  supposition. 

FIG.  213.  Leafy  branch  of  Uvularia  perfoliata. 

FIG.  214.  Leafy  and  flowering  branch  of  Baptisia  perfoliata. 

FIG.  215.  Lonicera  flava,  a  wild  Honeysuckle,  counate-perfoliate  as  to  the  upper 
leaves. 


108  MORPHOLOGY   OF   LEAVES. 

212.  "When  leaves  are  opposite,  the  perfoliation  (such  as  that 
of  Honeysuckles,  Fig.  215)  is  obviously  the  result  of  a  congeni- 
tal union  of  the  bases  of  the  pair  by  their  contiguous  edges. 
Leaves  connate  in  this  way  by  narrow  bases  are  not  rare  nor 
remarkable  ;  but  when  the  two  are  thus  coalescent  into  one  broad 
foliaceous  body,  giving  this  appearance  of  perfoliation,  the  term 
connate-perfoliate  is  used  to  express  it. 

213.  Vertical  Leaves,  those  with  blades  of  the  ordinary  kind, 
but  presenting  their  edges  instead  of  their  faces 

to  the  earth  and  sky,  or  when  erect  with  one 
edge  directed  to  the  stem  and  the  other  away 
from  it,  are  not  uncommon.  They  prevail  in 
the  Australian  Myrtaceae,  &c.,  and  occur  with 
less  constancy  in  the  Californian  Manzanitas, 
and  in  a  great  variety  of  herbs  and  shrubs.  The 
anomaly  involves  no  exception  to  the  rule  that  a 
leaf-blade  is  always  expanded  in  the  horizontal 
plane,  when  expanded  at  all ;  for,  except  in  equi- 
tant  leaves,  it  is  the  result  of  a  twist  of  the  petiole 
or  of  the  blade  itself.1  In  strongly  marked 
cases,  or  in  most  of  them,  the  organization  of 
the  epidermis  and  superficial  parenchyma  and 
the  distribution  of  the  stomata  are  the  same  on 
both  faces. 

214.  Eqnitant  Leaves  are  vertical  on  a  different 
plan.     They  are  conduplicate,  i.  e.  are   folded 


217 

together  lengthwise  on  their  middle,  the  upper  surface  thus  con- 
cealed within,  the  outer  alone  presented  to  the  air  and  light. 

1  Silphium  laciniatum,  the  so-called  Compass  Plant,  and  (hardly  less  so) 
S.  terebinthinaceum,  are  good  instances  of  the  kind,  most  of  the  leaves 
making  a  half-twist,  the  radical  ones  by  their  long  petioles.  In  the  former 
species,  the  pinnately  parted  blade  occasionally  makes  a  farther  twist,  so  as 
to  bring  the  upper  part  into  a  plane  at  right  angles  to  the  lower.  The 
blades  place  themselves  in  various  directions  as  respects  the  cardinal  points ; 
but  on  the  prairies  the  greater  number  affect  a  north  and  south  direction  of 
their  edges, — a  peculiarity  first  pointed  out.  in  the  year  1842,  by  General 
B.  Alvord,  U.  S.  JL 

FIG.  216.    Equitant  erect  leaves  of  Iris,  with  the  rootstock. 

FIG.  217.    A  section  across  these  leaves  at  the  base,  showing  the  equitant  character. 


THEIR    STRUCTURE   AND   FORMS. 


109 


Being  two-ranked  and  closely  crowded,  the  outer  ones  at  their 
base  fold  over  or  bestride  the  inner  (as  shown  in  the  sectional 
diagram,  Fig.  217),  whence  the  name  of  equitant.  Above,  the 
contiguous  halves  of  the  inner  face  conge nitally  cohere,  and  so 
produce  the  sword-shaped  or  linear  vertical  blade  which  is 
characteristic  of  Iris  (Fig.  216)  and  the  Iris  family.  In  most 
there  is  a  farther  complication,  of  an  excep- 
tional kind,  viz.  the  development  backwards 
of  a  portion  of  blade  from  the  midrib,  often 
forming  most  of  the  upper  part  of  such  leaves, 
which  therefore  may  really  be  said  to  develop 
in  the  vertical  plane. 

215.  Leaves  with  no  distinction  of  Parts,  *.  e. 
of  blade  and  petiole.     This  is  the  case  in  Iris 
(Fig.  216),  Daffodil,  the  Onion,  and  perhaps 
of  most  parallel-veined   leaves  of  Endogens. 
Those  expanded  in  the  horizontal  plane  may 
however  be  regarded  as  sessile  blades :  those 
which  are  not  expanded,  but  filiform,  or  needle- 
shaped  (acicular),  or  awl-shaped  (subulate),  may 
be  regarded  either  as  homologous  with  petioles, 
or  as  unexpanded  blades,  which  amounts  nearly 

to  the  same  thing  where  there  is  no  trace  of  a  petiole  at  base. 

Under  this  head  may  be  ranked  the  leaves  of  Pines  (Fig.  248)  ; 

also  both   the   subulate   and  the 

scale-shaped    and    adnate    leaves 

of  Arbor  Vitae,  Red  Cedar  (Juni- 

perus  Virginiana) ,  and  other  trees  < 

of  the  Cypress  tribe.  (Fig.  218.) 

216.  Stipules  serving  for  Blade. 
Lathyrus   Aphaca  is  a  good   in- 
stance of  this    (Fig.    219)  ;    the 
petiole    becoming    a    tendril,    the 
leaflets  which   its   relatives    bear 
being  wholly  wanting,  the  ample 
foliaceous     stipules    assume    the 
appearance   of  leaves.     In  some 

other  species  of  Lathyrus,  and  in  the  Pea,  equalty  large  stipules 
share  with  the  pair  or  pairs  of  leaflets  in  the  functions  of  foliage. 
On  morphological  evidence,  we  judge  that  the  singular  leaves  of 

FIG.  218.    A  twig  of  Arbor  Vitae,  with  both  awl-shaped  and  scale-shaped  leaves. 

FIG.  219.  Lathyrus  Aphaca:  portion  of  stem,  bearing  a  single  leaf,  which  consists 
of  a  pair  of  foliaceous  stipules,  and  a  petiole  in  the  form  of  a  tendril;  in  its  axil  a 
flower-stalk. 


110  MORPHOLOGY   OF  LEAVES. 

Baptisia  perfoliata,  shown  in  Fig.  214,  are  not  simple  blades, 
but  each  a  pair  of  stipules,  with  or  without  a  terminal  leaflet,  all 
completely  confluent  into  one  body.  The  related  species  of  the 
genus  have  trifoliolate  leaves  and  foliaceous  stipules  ;  hence  these 
simple  leaves  without  stipules  are  best  explained  in  this  way. 

217.  Phyllodia,  or  Petioles  serving  for  Blade.     Sometimes  the 
petiole  develops  foliaceous  margins,  or  wings,  as  in  the  Bitter 
Orange  and  in  Rhus  copallina.     These  are  efficient  as  foliage  in 
proportion  to  their  size.    These  are  not  to  be  confounded  with  the 
case  in  which  a  petiole  specially  develops  as  a  blade-like  organ, 
which  usurps  the  office  of  foliage.     A  petiole-blade  of  this  kind 
is  named  a  PHYLLODIUM.     Occurring  only  in  Exogens,  phyllodia 
are  generally  distinguished  from  true   blades  by   the   parallel 
venation,  and  always  by  their  normally  vertical  dilatation  ;  i.  e. 
they,  without  a  twist,  present  their  edges  instead  of  their  faces  to 
the  earth  and  sky.    The  common  and  most  familiar  phyllodia  are 
those  of  Acacias  in  Australia  (Fig.  223,  224),  where  they  form 
the  adult  foliage  of  over  270  out  of  less  than  300  species.     The 
true  lamina  of  these  is  bipinnate.     It  appears  on  seedlings,  and 
occasionally  on  later  growths.    Several  South  American  species  of 
Oxalis  produce  phyllodia.    So  likewise  do  our  tubular  or  trumpet- 
leaved  species  of  Sarracenia  in  that  portion  of  the  foliage  which 
develops   the   pitcher  imperfectly,  or  not  at   all.     Indeed,  all 
Sarracenia-leaves  are  phyllodia  with  the  back  in  most  of  them 
hollowed  out  into  a  tube  or  pitcher ;    and  the  terminal  hood 
answers  to  the  blade. 

§  3.     LEAVES  SERVING  SPECIAL  OFFICES. 

218.  Leaves  may  serve  at  the  same  time  both  their  ordinary 
and  some  special  use,  or  even  more  than  one  special  use.     For 
example,  in  Nepenthes   (Fig.   222)   there  is  a  well-developed 
blade,  usually  sessile,  which  serves  for  foliage,  a  prolongation  of 
its  tip  into  a  tendril,  which  serves  for  climbing,  then  an  extraor- 
dinary dilatation  and  hollowing  of  the  apex  of  this  into  a  pitcher 
for  a  very  special  use,  and  a  peculiar  development  of  the  apex 
of  this  into  a  lid,  closing  the  orifice  during  growth.     Among  the 
special  purposes  which  leaves  subserve,  and  the  study  of  which 
connects  singularities  of  morphology  with  teleology,  the  most 
remarkable  is  that  of 

219.  Leaves   specialised  for  the  Utilization  of  Animal  Matter. 
This  occurs  in  leaves  which  also  assimilate,  or  do  the  ordinary 
work  of  vegetation  ;  and  the  special  function  is  usually  taken  up 
by  some  particular  portion  of  the  organ.    The  details  of  this 


LEAVES   SERVING   SPECIAL  OFFICES. 


Ill 


subject  —  which  has  of  late  become  highly  interesting  —  belong 
to  physiology,  and  therefore  to  the  following  volume,  to  which 


all  historical  references  are  relegated.     Only  the  morphology  of 
such  leaves  is  here  under  consideration. 

220.  As  Ascidia  or  Pitchers,  vessels  for  maceration,  &c.  These 
occur  in  several  widely  different 
families  of  plants.  The  commonest 
are  those  of  the  Sarracenias,  natives 
of  Atlantic  North  America.  They 
are  evidently  phyllodia  (217),  the 
cavity  being  a  hollowed  dorsal  por- 
tion :  the  wing-like  or  foliaceous  por- 
tion, always  conspicuous  and  forming 
the  ventral  border,  makes  the  whole 
organ  or  most  of  it  in  the  earlier 
leaves  of  the  tubular  species.  The 
pitchers  of  S.  purpurea  (Fig.  221, 
225) ,  the  onty  species  which  extends 
north  of  Virginia,  are  open  cups, 
half  filled  with  water,  much  of  which 
may  be  rain,  in  which  abundance  of 
insects  are  usually  undergoing  macer- 
ation. In  S.  variolaris  (Fig.  226),  the  hooded  summit,  answer- 
ing to  the  blade  of  the  leaf,  arches  over  the  mouth  in  such  wise 


FIG.  220.  Pitchers  of  Heliamphora;  221.  of  Sarracenia  purpurea ;  222.  of  Nepenthes. 
223.  A  phyllodium  of  a  New  Holland  Acacia.  224.  The  same,  bearing  a  reduced  com- 
pound  blade. 

FIG.  225.  Pitcher-leaves  of  Sarracenia  purpurea;  one  of  them  with  the  upper  part 
oat  away. 


112  MORPHOLOGY   OF   LEAVES. 

as  to  mostly  exclude  the  rain ;  in  S.  psittacina  (Fig.  227)  the 

inflexed  and  inflated  hood  completely  excludes  it.  The  water 
which  these  contain  is  undoubtedly  a  secretion. 
All  entrap  flies,  ants,  and  various  insects,  which 
in  most  species  are  lured  into  the  pitcher  by  a 
sweetish  secretion  around  or  at  some  part  of  the 
orifice.1  Few  that  have  entered  ever  escape ; 
most  are  decomposed  at  the  bottom  of  the  cavity. 
In  Darlingtonia  Californica  (Fig.  228) ,  the  Cali- 
fornian  representative  of  Sarracenia,  the  inflated 
hood  guards  against  all  access  of  rain,  while  the 
orifice  is  freely  open  to  flying  insects  from  be- 
neath ;  and  a  singular  two-forked  appendage,  like 
to  a  fish-tail  (probably  the  homologUe  of  the 
blade) ,  overhangs  the  front.  The  inner  face  of 
this  appendage  is  besmeared  with  the  sweet  and 
viscid  secretion  which  allures  insects  to  the  open- 
ing. In  this  and  in  Sarracenia  variolaris,  the 
sweet  secretion  in  the  early  season  is  continued 
upon  the  edge  of  the  wing,  forming  a  saccharine 
trail  which  leads  from  near  the  ground  up  to  the 
orifice  of  the  pitcher.2  Fig.  220  represents  pitchers 
of  Heliamphora,  a  little-known  South  American 
representative  of  Sarracenia.  Its  wing  is  narrow 
and  inconspicuous,  the  mouth  widely  open  and 
directed  upward,  and  the  hood  reduced  to  a 
minute  and  upright,  probably  functionless  ap- 
pendage. In  Cephalotus — an  anomalous  plant 

of  Australia,  of  uncertain  affi- 
nity —  the  leaves  for  foliage 

are  dilated  phyllodia ;  among 

them    are   others    completely 

transformed  into  stalked  and 

short  pitchers,  with  thickened 

rim    and    a    well-fitting    lid, 

hinged  by  one  edge,  Fig.  229. 

The  particular  morphology  of  the  parts  is  not  well  made  out. 

1  This  sweet  secretion,  which  at  times  is  very  obvious  in  the  southern 
species,  has  also  been  detected  by  Mr.  Edward  Burgess  in  S.  purpurea ;  but 
it  is  rarely  seen,  and  probably  plays  no  important  part  in  the  capture  and 
drowning  of  the  multitude  of  insects  which  these  pitchers  are  apt  to  contain. 

2  This  trail  was  discovered  by  Dr.  J.  H.  Mellichamp,  of  South  Carolina. 
See  Proc.  Am.  Association  for  Advancement  of  Science,  xxiii.  113  (1871). 

FIG.  226.    Pitcher  of  Sarracenia  variolaria.    227.  Same  of  S.  psittacina. 


LEAVES   SERVING   SPECIAL  OFFICES.  113 

221.  The  pitcher-bearing  leaf  in  Nepenthes  has  been  referred 
to  (218,  Fig.  222)  :  of  this  there  are  various  species,  all  of  them 
somewhat  woody  climbing 

plants  of  tropical  Asiatic 
and  African  islands  of 
the  southern  hemisphere, 
some  of  them  familiar  in 
conservatory  cultivation. 
Here  the  tendril  may  be 
regarded  as  a  prolonged 
extension  of  the  midrib 
of  the  blade,  and  the 
pitcher,  with  its  hinged 

lid,  as  a  peculiar  development  from  its  apex.  The 
water  contained  in  the  pitcher  is  a  secretion,  much 
of  which  appears  before  the  lid  opens  ;  and  a  sweetish 
excretion  at  the  orifice  lures  insects.  The  presence 
of  these  in  the  pitcher  increases  the  watery  secre- 
tion in  which  the  animals  are  drowned ;  and  this 
secretion  is  ascertained  to  have  a  certain  digestive 
power.1 

222.  The  aquatic  sacs  of  Utricularia  or  Bladder- 
wort  are  diminutive  ascidia,  always  under  water, 

and  with  lid  opening  inward,  like  a  valve,  preventing  the  exit 
of  minute  animals  entrapped  therein.2  Morphologically,  they 
are  doubtless  leaves  or  parts  of  leaves. 

223.  As   Sensitive  Fly-traps.      The  leaves  of  all   species   of 
Drosera  or  Sundew  are  beset  with  stout  bristles  tipped  with  a 
gland,  which  secretes  and  when  in  good  condition  is  covered 
by  a  drop  of  a  transparent  and  very  glairy  liquid,  sufficiently 
tenacious  to  hold  fast  a  fly  or  other  small  insect.     Adjacent 
bristles,  even  if  not  touched,  in  a  short  time  bend  towards  those 
upon  which  the  insect  rests,  and  thus  bring  their  glands  also 
into  contact  with  it.     In  Drosera  filifonnis,  the  leaves  are  fili- 
form, with  no  distinction  of  petiole  and  blade.     In  D.  rotundifolia 
and  other  common  round-leaved  species,  there  is  a  clear  distinc- 

1  This  was  first  made  out  by  J.  D.  Hooker,  and  announced  in  his  address, 
as  President  of  the  British  Association  for  the  Advancement  of  Science,  at 
Edinburgh,  1871. 

2  Darwin,  Insectivorous  Plants,  395.     Cohn,  Beitrage  zur  Biologie  der 
Pflanzen,  1875.     Mrs.  Treat,  in  The  Tribune,  New  York,  September,  1874, 
and  Card.  Chron.  1875,  303. 

FIG.  228.    Pitcher  of  Darlingtonia  California.    229.  Pitcher  of  Cephalotua  follicu- 
laria,  with  lid  open. 


114  MORPHOLOGY   OF   LEAVES. 

tion  of  petiole  and  blade,  and  the  stalked  glands  thickly  beset 
the  whole  upper  surface  of  the  latter.  A  small  insect  alighting 
thereon  is  helpless,  and  is  soon  touched  by  all  the  glands  within 
reaching  distance ;  also  the  blade  itself  commonly  incurves, 
taking  part  in  the  general  movement.  It  has  recently  been 
demonstrated  that  the  captured  insect  is  fed  upon,  and  that  the 
plant  thereby  receives  nourishment.  Here  leaves  which  do  the 
normal  assimilative  work  of  vegetation,  but  somewhat  feebl}- 
(having  a  comparatively  small  amount  of  chlorophyll),  have 


also  the  power  and  the  habit  of  obtaining  ready-organized  food 
by  capture,  and  are  benefited  by  it. 

224.  Species  of  Drosera  inhabit  most  parts  of  the  world,  and 
the  genus  is  numerous  in  species.  A  near  relative,  Dionaea,  is 
of  a  single  species,  D.  muscipula  (Venus's  Fly-trap) ,  inhabiting 
only  a  limited  district  in  the  sandy  eastern  border  of  North 
Carolina.  It  is  more  strikingly  sensitive  and  equally  carnivo- 
rous, but  in  a  different  way.  It  is  destitute  of  stalked  and  viscid 
glands.  The  apparatus  for  capture  and  digestion  is  the  two- 
valved  body  at  the  top  of  each  leaf.  (Fig.  230,  231.)  If  this 

FIG.  230.  A  plant  of  Dionaea  muscipula,  reduced  in  size.  231  Three  of  the  leaves, 
of  almost  the  natural  size ;  one  of  them  open,  the  others  closed.  Probably  a  fly  is  never 
caught  by  the  teeth,  in  the  manner  here  represented. 


LEAVES   SERVING   SPECIAL   OFFICES.  115 

be  taken  for  the  leaf-blade,  the  part  below  would  be  a  broadly- 
winged  foliaceous  petiole.  If  the  latter  be  the  true  blade,  the 
apparatus  in  question  must  be  reckoned  as  a  peculiar  terminal 
appendage.  Both  are  moderately  green,  and  act  as  foliage. 
The  specially  endowed  terminal  portion  acts  also  in  a  decidedly 
animal-like  manner.  When  either  of  the  three  or  four  slender 
bristles  of  the  upper  surface  are  touched,  the  trap  suddenly 
closes,  by  a  movement  ordinarily  quick  enough  to  enclose  and 
retain  a  fly  or  other  small  insect.  The  intercrossing  of  the  stout 
marginal  bristles  detains  the  captive,  unless  it  happens  to  be 
small  enough  to  escape  by  the  intervening  little  openings. 
Otherwise,  the  sides  soon  flatten  and  are  brought  firmly  into 
contact,  and  a  glairy  secretion  is  poured  out  from  numerous 
immersed  glands  :  this,  with  the  extracted  juices  of  the  macerated 
insect,  is  after  some  time  reabsorbed ;  the  trap,  if  in  a  healthy 
condition,  now  re-opens  and  is  ready  for  another  capture.  For 
references  to  the  now  copious  literature  of  this  whole  subject,  and 
for  its  physiological  treatment,  the  succeeding  volume  should  be 
consulted. 

225.  Leaves  for  Storage.  Nutritive  matter  is  stored  in  leaves 
in  many  cases,  and  not  rarel}'  in  leaves  which  at  the 
same  time  are  subserving  the  purpose  of  foliage. 
This  occurs  in  all  fleshy  leaves,  to  a  greater  or  less 
extent,  according  to  the  degree  of  thickening  or 
accumulation.  The  leaves  of  the  Century  Plant 
or  Agave,  for  instance,  are  green  and  foliaceously 
efficient  at  the  surface,  while  the  whole  interior  is 
a  store-house  of  farinaceous  and  other  nutritious 
matter,  as  much  so  as  is  a  potato.  The  leaves 
of  various  species  of  Aloe,  Mesembryanthemum, 
Sedum,  and  other  "succulent"  plants  (in  which 
a  large  part  of  the  accumulation  is  water)  are  not 
rarely  so  obese  as  to  lose  or  much  disguise  the 
foliaceous  appearance.  Sometimes  one  portion  of 
a  leaf  is  of  normal  texture  and  use,  while  another 
is  used  as  a  reservoir  for  the  nourishment  which  the 
foliaceous  part  has  produced.  Fig.  232,  a  leaf  from 
the  bulb  of  White  Lily,  the  base  of  which  forms 
one  of  the  bulb-scales,  is  an  instance  of  the  kind.1  The  most 

1  In  Dicentra  Cucullaria  and  (more  strikingly  from  the  sparseness  of  the 
grains)  in  D.  Canadensis,  the  matter  elaborated  in  the  much  dissected  blade  is 
conveyed  to  the  very  base  of  the  long  petiole,  and  there  deposited  in  a  con- 

FIG.  232.  A  radical  leaf  of  the  White  Lily,  with  Its  base  thickened  into  a  bulb-scale, 
which  is  cut  across  to  show  its  thickness. 


116  MORPHOLOGY  OF  LEAVES. 

decisive  instance  of  leaves  used  for  storage  of  food  is  in  that 
material  provision  for  the  nourishment  of  the  embryo  in  germU 
nation,  in  which  the  first  leaves,  the  cotyledons,  are  turned  to  this 
account.  (21-37,  &c.)  After  or 
while  discharging  this  special  duty, 
the  cotyledons  may  fulfil  their  gen- 
eral office,  by  serving  as  foliage  (as 
in  Maples,  Fig.  8,  and  Pumpkins, 
Fig.  47)  ;  or,  through  various  inter- 
mediate conditions,  they  may  be 
wholly  devoted  to  storage,  as  in 
'the  Pea,  Oak,  Horsechestnut,  &c. 
(Fig.  37-43.) 

226.  Leaves  as  Bulb-scales,  how- 
ever, are  for  the  most  part  wholty 
applied  to  this  use,  being  leaves 
reduced    to    short     scales    or    to 

1  concentric  coats,  and  thickened 
throughout  by  nutritive  deposit. 
The  accumulation  of  such  leaves 
forms  the  mass  of  the  bulb,  as  of 
the  Lily,  Fig.  118,  Onion,  Fig. 
113,  &c.,  also  of  bulblets.  (120.) 

227.  Leaves  as  Bud-scales,  being 
for  protection  of  nascent  parts,  have 
been  explained  under  buds.   (70.) 
The  evidence  of  foliar  nature  af- 
forded by  transition  is  well  exhib- 
ited by  the  Sweet  Bucke3'e,  although 
the  whole  series  of  gradations,  from 
bud-scales    to    compound    leaves, 
is  seldom  seen  united  in  one  bud, 
as  in  Fig.  233.     In  this  case,  the 
bud-scales    are    homologous    with 
petioles.  In  Magnolia,  they  consist 
of  stipules   (Fig.  81,  82)  :  in  the 

Lilac,  they  are  homologous  with  leaf-blades.  The  two  pairs  of 
bud-scales  which  subtend  and  protect  through  winter  the  nascent 
head  of  flowers  of  Cornus  florida  are  morphologically  the  apex  of 

centrated  condition,  in  the  form  of  a  solid  grain,  which  remains  for  next 
year's  use,  the  whole  leaf  except  this  thickened  base  dying  away  at  the  close 
of  the  short  season's  growth. 

FIG.  233.  Leaves  of  a  developing  bud  of  the  Low  Sweet  Buckeye  (.Escnlus  parvi- 
flora),  showing  a  nearly  complete  set  of  gradations  from  a  scale  to  a  compound  leaf  o» 
five  leaflet* 


LEAVES   SERVING   SPECIAL   OFFICES.  117 

blades.    When  the  blossoms  develop  in  spring,  these  scales  grow 
from  beneath,  greatly  expand,  and  become  obovate  or  obcordate 
petaloid  leaves,  the  brown  terminal  notch 
of  which  is  the  bud-scale,  which  was  un- 
able to  take  part  in  the  vernal  growth. 

227°.  Leaves  as  Spines.  All  gradations 
may  be  found  between  spiny-toothed  leaves 
(as  in  Holly),  in  which  teeth  are  pointed  and 
indurated,  and  leaves  which  are  completely 
contracted  into  a  simple  or  multiple  spine. 
Indeed,  such  a  transition  is  seen  in  the  Bar- 
berry, Fig.  234.  The  foliar  nature  of  such 
spines  is  manifest  from  their  position,  sub- 
tending a  bud  from  which  the  foliage  of  the 
season  proceeds,  and  themselves  not  sub- 
tended by  any  organ.  In  some  Astragali, 
the  petiole  of  a  pinnate  leaf  indurates  into 
a  slender  spine  and  persists,  the  leaflets 
early  falling.  The  spine  in  Fouquiera  is  a 
portion  of  the  lower  side  of  the  petiole  or 
midrib,  indurated  and  persistent,  the  rest 
of  the  leaf  separating  by  splitting  when  it 
has  served  its  office.1 

228.  Leaves  adapted  to  Climbing.  Some  plants  climb  by  the 
action  of  the  stem  or  of  certain  branches  specially  adapted  to 
this  purpose  (99)  :  others  gain  the  needful 
support  by  means  of  their  leaves  (101)  ;  some- 
times by  an  incurvation  of  the  tips,  either  of 
a  simple  blade  as  in  Gloriosa,  or  small  partial 
blades,  as  inAdlumia,  and  often  in  Clematis, 
thereby  grappling  the  support ;  sometimes  by 
the  petiole  making  a  turn  or  two  around  a 
support  (as  in  Maurandia,  climbing  Antirrhi- 
nums, Rhodochiton,  and  Solanum  jasminoides,  | 
Fig.  235)  ;  sometimes  by  the  transformation 
of  one  or  more  leaflets  of  a  compound  leaf 
into  tendrils,  as  in  the  Pea  and  Vetch  (Fig. 
204)  ;  sometimes  by  the  suppression  of  all  the 
leaflets  and  the  conversion  of  the  whole  petiole  into  a  tendril,  as 
in  Lathyrus  Aphaca  (Fig.  219)  ;  and  perhaps  by  the  conversion 

1  Described  in  Plantse  Wrightianse,  ii.  63. 

FIG.  234.    A  vernal  shoot  of  common  Barberry,  showing  a  lower  leaf  in  the  normal 
state ;  the  next  partially,  those  still  higher  completely,  transformed  into  spines. 

FIG.  235.  Solanum  jasminoides,  climbing  by  coiling  and  at  length  indurating  petioles. 


118  MORPHOLOGY   OP   LEAVES. 

of  a  pair  of  stipules  into  tendrils  in  Smilax.  At  least  the  ten- 
drils here  occupy  the  position  of  adnate  stipules.  The  tendrils 
of  Cucurbitaceae  are  peculiar  and  ambiguous,  on  account  of  their 
lateral  and  extra-axillary  position  and  the  manner  in  which  the 
compound  ones  develop  their  branches.  But  they  are  doubtless 
partly  if  not  wholly  foliar.1 

229.  Petaloid  Leaves,  Bracts.     Certain  leaves,  situated  near  to 
flowers,  and  developing  little  or  no  chlorophyll  in  their  paren- 
chyma, exchange  the  ordinary  green  hue  and  herbaceous  texture 
for  the  brighter  colors  and  more  delicate  structure  which  are 
commonly  seen   in  and  thought  to  characterize  flower-leaves. 
Such  are  said  to  be  colored,  meaning,  as  applied  to  foliage,  of 
some  other  color  than  green.     As  petals  are  the  type  of  such 
colored  parts,  they  are  said  to  be  petaloid,  i.  e.  petal-like.     They 
are  like  petals,  moreover,  in  one  of  the  purposes  which  these  sub- 
serve. (299.)     Examples  of  these  petaloid  leaves  are  seen  in  the 
shrubby  Mexican  Euphorbia  called  Poinsettia,  in  Salvia  splen- 
dens,  most  species  of  Castilleia  or  Painted  Cup,   also  in  the 
white  hood  of  Calla  and  Richardia  JEthiopica  (called  Calla  Lily), 
and  in  the  four  white  leaves  which  subtend  the  flower-head  of 
Cornus  florida,  and  of  the  low  herbaceous  Cornel,  C.  Canaden- 
sis.  (Fig.  294.)    Such  leaves,  being  in  proximity  to  flowers,  and 
all  others  which  are  within  a  flower-cluster  or  are  borne  by 
flower-stalks,  receive  the  special  name  of  BRACTS.     More  usually 
bracts  are  not  petaloid,  but  different  in  size  or  shape  from  ordi- 
nary leaves,  either  by  abrupt  change  or  gradual  transition.    Not 
uncommonly  they  are  reduced  to  scales  or  mere  rudiments  or 
vestiges  of  leaves,  of  no  functional  importance. 

230.  Flower-Leaves.     The  morphology  of  leaves  extends  not 
only  to  "  the  leaves  of  the  blossom,"  more  or  less  accounted  as 
such  in  common  parlance,  but  also  to  its  peculiar  and  essential 
organs,  the  relation  of  which  to  leaves  is  more  recondite.     Their 
morpholog}"  needs  to  be  treated  separately,  and  to  be  preceded 
by  a  study  of  the  arrangement  of  leaves  and  of  blossoms. 

1  The  most  satisfactory  interpretation  may  be  that  of  Braun  and  Wydler, 
adopted  by  Eichler  (Bliithendiagramme,  i.  304) :  that  the  flower  of  Cucur- 
bita  and  its  peduncle  represent  the  axillary  branch,  the  tendril  by  its  side 
answers  to  one  of  the  bractlets  (that  of  the  other  side  being  suppressed), 
and  the  supernumerary  branch  springs  from  the  axil  of  the  tendril.  This 
makes  of  the  tendril  a  simple  leaf,  of  which  the  branches  are  the  ribs.  But 
the  tendril-divisions  are  evidently  developed  in  spiral  order,  and  in  vigorous 
growths  occupy  different  heights  on  the  tendril-axis.  This  favors  Naudin'a 
view,  that  the  main  tendril  is  cauline,  and  its  divisions  leaves. 


PHYLLOTAXY,  OR   LEAF-ARRANGEMENT.  119 

CHAPTER  IV. 

PHYLLOTAXY,  OR  LEAF-ARRANGEMENT. 

SECTION  I.     THE  DISTRIBUTION  OF  LEAVES  ON  THE  STEM. 

231.  PHYLLOTAXY  (or  PhyUotaxis)  is  the  study  of  the  distri- 
bution of  leaves  upon  the  stem  and  of  the  laws  which  govern  it. 
The  general  conclusion  reached  is,  that  leaves 

are  distributed  in  a  manner  to  economize  space 
and  have  a  good  exposure  to  light,  &c.,  and 
that  this  economy  on  the  whole  results  from 
the  formation  of  leaves  in  the  bud  over  the 
widest  intervals  between  the 
leaves  next  below.1   Leaves 
are  arranged  in  a  consider- 
able variet}7  of  ways,  which 
all  fall  under  two  modes,  the 
Verticillate    and    the    Alter- 
nate (13),  but   which  may 
also  be  termed  the  Cyclical  ( 
and  the  Spiral. 

232.  Alternate  leaves  are 
those   which   stand   singly, 
one  after  another ;  that  is, 
with  one  leaf  to  each  node 

or  borne 

on  one  height  of  stem.  Verticillate  leaves  are 
those  with  two  or  more  at  the  same  height  of/ 
stem,  circularly  encompassing  it,  i.  e.  forming! 
a  Verticil  or  Whorl.  Verticillate  and  whorled 
are  synonymous  terms  to  denote  this  arrange- 
ment. These  two  kinds  of  leaf-arrangement 
are  commonly  ranked  as  three,  viz.  alternate, 
opposite,  and  whorled.  But  the  opposite  is 
only  the  simplest  case  of  the  whorled,  being 

1  For  the  most  comprehensive  discussion  of  phyllotaxy  in  connection  with 
development,  and  in  view  of  these  relations,  see  Hofmeister,  Allgemeine 
Morphologie,  §  11,  and  Chauncey  Wright,  Mem.  Amer.  Academy,  ix.  389. 
FIG.  236,  Alternate,  237,  Opposite,  238,  Verticillate  or  whorled  leaves. 


120  PHYLLOTAXY,    OR    LEAF-ARRANGEMENT. 

that  in  which  the  members  are  reduced  to  two.  This  case  is 
so  much  commoner  than  whorls  of  three  and  of  higher  numbers 
that  it  took  from  the  first  its  special  name  of  opposite,  so  that  in 
descriptions  the  phrase  "  leaves  verticillate  "  implies  more  than 
two  leaves  in  the  whorl.  But  it  should  be  kept  in  mind  that 
"leaves  opposite  "  is  the  same  as  "  leaves  in  whorls  of  two." 

233.  The  greater  number  of  phaenogamous  plants  (all  but  the 
monocotyledonous  class)  begin  with  verticillate  leaves,  mostly 
of  the  simplest  kind  (z.  e.  cotyledons  opposite)  :  some  continue 
verticillate  throughout ;  some  change  in  the  first  leaves  of  the 
plumule  or  after  the   first  pair   into  alternate,  and   again  into 
verticillate  in  or  toward  the  blossom,  in  the  interior  of  which  the 
alternate  arrangement  may  be  again  resumed.    As  Nature  passes 
»~;adily  from  the  one  mode  to  the  other  on  the  same  axis,  we 
may  expect  that  the  two  may  be  comprised  under  some  common 
expression.     But  they  have  not  yet  been  combined,  except  by 
gratuitous   or   somewhat  forced   hypotheses ;    so   that   for  the 
present  they  should  be  treated  in  morphology  as  primarily  dis- 
tinct arrangements.1 

234.  Verticillate   or  Cyclical  Arrangement.      Here  the  leaves 
occupy  a  succession  of  circles,  or  form  whorls  around  the  stem, 
two,  three,  four,  five,  &c.,  in  each  whorl.     According  to  the 
number,  the  leaves  are  opposite,  ternate,  quaternate,  quinate,  and 
so  on.     The  characteristic  of  the  individual  whorl  is  that  the 
members  stand  as  far  apart  from  each  other  as  then-  number 
renders  possible,  i.  e.  they  divide  the  circle  equally.    Thus,  when 
only  two,  or  opposite,  their  midribs  or  axes  of  insertion  have  an 
angular  divergence  (as  it  is  termed)  of  180° ;  when  three,  of 
120°  ;  when  four,  90°  ;  when  five,  72°. 

235.  The  characteristic  of  the  whorls  in  relation  to  each  other 
is,  that  the  members  of  successive  whorls  stand  over  or  under 
the  intervals  of  the  adjacent  ones.     In  other  words,  successive 
whorls  alternate  or  decussate.     This  economizes  space  and  light, 
or  gives  the  best  distribution  which  the  cyclical  sj-stem  is  capa- 
ble of.     And  it  is  in  accordance  with  the  general  conclusion  of 
Hofmeister's  investigation  of  the  origin  of  phyllotaxic  arrange- 
ments in  the  nascent  bud,  viz.  that  new  members  originate  just 
over  the  widest  intervals  between  their  predecessors  next  below. 
Thus,  in  opposite  leaves  or  whorls  of  two  (Fig.  237),  the  suc- 
cessive  pairs  decussate   or  cross  at  right  angles,  and  so  four 

1  It  is  readily  seen  that  whorls  may  be  produced  by  the  non-development 
of  the  internodes  between  the  leaves  of  a  series  of  two,  three,  five,  or  more 
in  alternate  order.  The  difficulty  is  that  the  members  of  the  next  whorl  do 
not  follow  the  order  that  they  should  upon  this  supposition. 


DISTRIBUTION  OF   LEAVES   ON   THE   STEM.  121 

straight  equidistant  vertical  ranks  are  produced.  In  ternate  or 
trimerous  whorls  there  are  six  vertical  ranks  ;  in  quaternate  or 
tetramerous  whorls,  eight  vertical  ranks,  and  so  on.1 

236.  The   cases   in   which   successive 
pairs  of  leaves  do  not  decussate  at  right 
angles,  or  the  members  of  whorls  are  not 
exactly  superposed  to  intervals,  but  as  it 
were  wind  spirally  (as  in  Dipsacus,  many 
Caryophyllaceae,  &c.) ,  may  some  of  them 
be  explained   by   torsion   of  the    stem, 
such  as  is  very  manifest  in  numerous  in- 
stances ;  and  others  may  be  resolved  into 

instances  of  alternate  leaves  simulating  or  passing  into  whorls 
by  the  non-development  of  internodes.2 

237.  Alternate  or  Spiral  Arrangement.     Here  the  leaves  are 
distributed  singly  at  different  heights  of  the  stem,  and  at  equal 
intervals  as  respects  angular  divergence.  (Fig.  236.)    This  angu- 
lar divergence  (i.  e.  the  angular  distance  of  any  two  successive 
leaves)  differs  in  the  various  kinds  of  this  system  of  phyllotaxy, 
but  is  alwa}-s  large  enough  to  place  the  leaves  which  immediately 

1  These  vertical  ranks  have,  by  some  German  botanists,  been  named 
Orthostichies ;  but  this  technical  Greek  is  no  clearer  and  no  shorter  than  the 
equivalent  English,  which  answers  every  purpose. 

2  In  Lilium  Canadense,  superbum,  &c.,  with  whorls  of  variable  number 
of  leaves  and  vague  relation  to  each  other  (when  of  the  same  number  some- 
times the  members  superposed),  and  above  and  below  passing  into  the  alter- 
nate arrangement  normal  to  the  family,  these  whorls  are  evidently  formed 
of  alternate  leaves  brought  together  by  non-development  of  internodes. 

Here  may  also  be  mentioned  the  not  uncommon  anomaly  in  Fir-cones, 
notably  those  of  Norway  Spruce,  the  normal  phyllotaxy  of  which  is  simply 
spiral,  but  in  occasional  instances  the  cone  is  composed  of  pairs  of  opposite 
scales,  spirally  arranged,  i.  e.  the  pairs  not  decussating  at  right  angles,  thus 
forming  double  spirals.  In  the  abnormal  spruce-cones,  the  fractions  usually 
observed  are  -f$  or  ^,  or,  as  expressed  by  Braun,  (|)^  and  (^)-^g-. 

Braun's  mode  of  notation  for  the  ordinary  succession  (i.  e.  the  decussation) 
of  opposite  leaves  is  (£)i,  the  |  meaning  that  the  two  leaves  of  the  pair  are 
half  the  circumference  of  the  circle  apart,  the  J  denoting  that  each  leaf  of 
the  succeeding  pair  diverges  one  fourth  of  the  circumference  from  the  "pre- 
ceding. Braun  finds  cases  in  which  pairs  (and  equally  whorls)  are  super- 
posed (e.g.  certain  species  of  Mesembryanthemum  and  Euphorbia),  these 
are  expressed  in  this  notation  by  the  formula  (|)£,  that  is,  the  corresponding 
leaves  of  the  succeeding  pair  diverge  180°  from  their  predecessors.  He 
recognizes  also  some  cases  of  intermediate  divergence;  such  as  (£)$  in  the 
upper  leaves  of  Mercurialis  perennis,  (i)^r  on  certain  stems  of  Linaria  vul- 
garis,  (i)y5$  exceptionally  in  the  leaves  of  Epilobium  angustifolium  and  the 
scales  of  Norway  Spruce,  (i)-^  exceptionally  in  the  scales  of  Norway 
Spruce.  See  Ordnung  des  Schuppen  an  der  Tannenzapfen,  376,  &c. 

FIG.  239.    Ground-plan  diagram  of  six  trimerous  whorls,  showing  their  alternation. 


122  PHYLLOTAXY,   OB   LEAF- ARRANGEMENT. 

follow  each  other  in  the  ascending  order  upon  different  sides  of 
the  axis  :  it  also  secures  an  advantageous  spacing  of  the  leaves 
over  the  whole  length  of  the  axis.  Their  vertical  distance  from 
each  other  of  course  depends  on  the  length  which  the  internodes 
attain,  which  is  a  matter  of  growth  and  is  very  variable  ;  but 
their  angular  distance  is  fixed  in  the  kind  or  numerical  plan  of 
the  particular  phyllotaxy,  and  is  uniform  throughout. 

238.  The  leaves  are  said  to  be  alternate,  because  they  come 
one  after  another,  now  on  this  side,  then  on  that,  as  they  ascend 
the  stem.     The  arrangement  is  said  to  be  spiral,  because  if  a  line 
be  drawn  or  a  thread  extended  from  the  base  or  insertion  of  one 
leaf  to  that  of  the  next  higher,  and  so  on,  taking  in  all  the  leaves, 
it  forms  a  helix,  more  or  less  loose  or  close  according  to  the 
development  of  the  internodes.  (See  Fig.  242.)    This  imagined 
spiral  line  ascends  continuously,  without  a  break ;  and  on  it  the 
leaves  are  equably  laid  down.1 

239.  Almost  all  the  ordinary  instances  of  spiral  phyllotaxy 
belong  to  one  series,  having  very  simple  arithmetical  relations. 
So  that  this  may  be  taken  as  the  type,  and  the  few  others  re- 
garded as  exceptions  or  sometimes  as  modifications  of  it.     The 
kinds  are  simply  designated  by  the  number  of  vertical  ranks  of 
leaves :    they  are  technically   named   by  prefixing   the   proper 
Greek  numeral  to  the  word  meaning  row  or  rank.     The  arrange- 
ment called 

Distichous,  or  Two-ranked,  is  the  simplest  and  among  the  com- 
monest, occurring,  as  it  does,  in  all  Grasses  and  many  other 
monocotyledonous  plants,  in  Linde&s,  Elms,  and  many  dico- 
tyledonous genera.  Here  the  leaves  are  disposed  alternately  on 
exactly  opposite  sides  of  the  stem  (as  in  Fig.  1)  ;  the  second 
leaf  being  the  farthest  possible  from  the  first,  as  is  the  third  from 
the  second  ;  the  third  therefore  over  the  first,  and  the  fourth  over 
the  second,  and  so  on,  thus  forming  two  vertical  ranks.  The 
angular  divergence  is  here  half  the  circumference,  or  180° ;  and 
the  phyllotaxy  may  be  represented  by  the  fraction  ^,  which  desig- 
nates the  angular  divergence,  while  its  denominator  expresses 
the  number  of  vertical  ranks  formed. 

Tristichous,  or  Three-ranked,  is  the  next  in  the  series,  and  is 

1  But  when  we  reach  a  leaf  which  stands  directly  over  a  lower  and  older 
one,  we  say  that  one  set  or  spire  is  completed,  and  that  this  leaf  is  the  first 
of  a  succeeding  set  or  spire.  From  analogy  of  such  an  open  spire  to  the 
closed  cycle  of  a  whorl  of  leaves,  it  is  not  unusual  to  designate  the  former 
likewise  as  a  cycle.  Yet  it  is  better  (with  Eichler)  to  restrict  that  term,  and 
the  adjective  cyclical,  to  verticillate  phyllotaxy,  or  to  whorls,  to  which  it 
properly  and  etymologically  belongs. 


DISTRIBUTION  OF   LEAVES   ON  THE  STEM. 


123 


less  common,  though  not  rare  in  monocotyledonous  plants.  Fig. 
240  illustrates  it  in  a  Sedge,  and  241  is  a  diagram  in  horizontal 
section,  as  of  a  bud ;  both 
extending  to  six  leaves  or 
two  turns  of  the  spiral.  The 
fraction  £  designates  this 
arrangement.  The  angular 
divergence,  or  distance  of 
the  axis  of  the  first  leaf 
from  the  second,  and  so  on, 
is  one  third  of  the  circum- 
ference (or  120°)  :  conse- 
quently the  fourth  leaf  comes 
over  the  first,  the  fifth  over 
the  second,  the  sixth  over 
the  third,  and  so  on ;  that 
is,  the  leaves  fall  into  three 
vertical  ranks.  The  spiral 
character  here  begins  to  be 
manifest,  or  becomes  so  by 
drawing  a  line  on  either  fig- 
ure from  the  axis  or  midrib 
of  the  first  leaf  to  that  of 
the  second,  and  so  on  to  the 
sixth,  forming  a  helix  of 
two  turns.1 

Pentastichous ,  or  Five-ranked,  sometimes  termed  the  quincuncial 
arrangement.  This  is  the  most  common  in  alternate-leaved 
dicotyledonous  plants.  It  is  shown  in  Fig.  236  (on  a  branch 
of  Apple-tree) ,  and  by  diagrams,  displaying  the  spiral  character, 
in  Fig.  242,  243.  The  angular  distance  from  the  first  to  the 
second  leaf  (passing  the  shorter  way)  is  §  of  the  circumference, 
or  144°.  But  the  spiral  line  makes  two  turns  round  the  stem, 
on  which  six  leaves  are  laid  down,  with  angular  divergence  of  f , 

1  The  line  is  supposed  to  follow  the  nearest  way,  and  the  divergence  is 
counted  as  £,  this  being  the  simplest  and  most  convenient.  If  for  any  reason 
the  longer  way  is  preferred,  then  the  angular  divergence  would  be  expressed 
by  the  fraction  |. 

FIG.  240.  Piece  of  a  stalk,  with  the  sheathing  bases  of  the  leaves,  of  a  Sedge-Grass 
(Carex  crus-corvi),  showing  the  three-ranked  arrangement.  241.  Diagram  of  the  cross- 
section  of  the  same.  The  leaves  are  numbered  in  succession. 

FIG.  242.  Diagram  of  position  of  six  leaves  in  the  five-ranked  arrangement :  a  spiral 
line  is  drawn  ascending  the  stem  and  passing  through  the  successive  scars  which  mark 
the  position  of  the  leaves  from  1  to  6.  It  is  made  a  dotted  line  where  it  passes  on  the 
opposite  side  of  the  stem,  and  the  scars  2  and  5,  which  fall  on  that  side,  are  made 
fainter.  243.  A  plane  horizontal  projection  of  the  same;  the  dotted  line  passing  from 
the  edge  of  the  first  leaf  to  the  second,  and  so  on  to  the  fifth  leaf,  which  completes  the 
turm ;  M  the  sixth  would  come  directly  before,  or  within,  the  first. 


124  PHYLLOTAXY,   OR   LEAF-ABE  ANGEMENT. 

and  the  sixth  is  the  first  to  come  over  any  one  below ;  the 
seventh  comes  over  the  second,  the  eighth  over  the  third,  &c. 
The  leaves  are  thus  brought  into  five  vertical  ranks  ;  but  these 
five  leaves  are  laid  down  on  two  turns  of  the 
helix  (the  sixth  beginning  the  second  revolu- 
tion) ;  the  angular  divergence  of  the  leaves  in 
order  is  f ,  or  144° ;  the  angular  distance  of 
the  vertical  ranks,  72°.  This  is  a  very  advan- 
tageous distribution  for  ordinary  foliage  on 
erect  or  ascending  branches.  Its  formula  is  f ,  expressing  the 
angular  divergence,  the  denominator  also  indicating  the  number 
of  vertical  ranks,  the  numer- 
ator indicating  the  number 
of  revolutions  made  in  add- 
ing one  leaf  to  each  rank. 
Fig.  244  illustrates  this  ar- 
rangement on  a  cone  of 
American  Larch,  the  scales 
of  which  are  homologous 
with  leaves,  the  numbers 
in  sight  are  affixed,  and  those  of  the  whole 
cone  displayed  on  a  plane  at  the  side. 

Octostichous,  or  Eight-ranked,  a  less  common 
arrangement,  occurs  in  the  Holly,  Aconite, 
the  radical  leaves  of  Plantago.  It  has  the 
angular  divergence  of  135°,  or  f  of  the  cir- 
cumference, and  the  leaves  in  eight  ranks, 
the  ninth  over  the  first  and  at  the  completion 
of  the  third  revolution  :  it  is  therefore  repre- 
sented by  the  fraction  f . 

240.  The  obvious  relations  of  the  fractions 
vi  %•>  $•>  t>  representing  the  primary  forms  of 
spiral  phyllotaxy,  are  that  the  sum  of  any  two 
numerators  is  the  numerator  of  the  next  suc- 
ceeding fraction,  and  the  same  is  true  of  the 
denominators  ;  also  the  numerator  is  the  same 
as  the  denominator  of  the  next  but  one  pro- 
ceeding fraction.  Following  these  indications, 
the  series  may  be  extended  to  ^,  ^,  £f ,  §J,  &c.  Now  these 

FIG.  244.  A  cone  of  the  small-fruited  American  Larch  (Larix  Americana),  with 
the  scales  numbered,  exhibiting  the  five-ranked  arrangement. 

FIG.  245.  An  offset  of  the  Houseleek,  exhibiting  the  5-13  arrangement;  the  leaves 
in  sight  numbered,  the  14th  over  the  first,  the  19th  over  the  6tb,  &c. 

FIG.  246.  Cone  of  White  Pine  (Pinus  Strobus)  with  scales  numbered  from  bottom, 
and  some  secondary  spirals  marked. 


DISTRIBUTION  OF   LEAVES   ON  THE  STEM.  125 

cases  actually  occur,  and  ordinarily  only  these.1  The  -^  and 
A  are  not  uncommon  in  foliage.  The  rosettes  of  the  House- 
leek  exhibit  the  ^  or  thirteen-ranked  arrangement,  as  also  does 
the  cone  of  Pinus  Strobus,  the  14th  leaf  falling  over  the  first. 
(Fig.  246.)  The  &  is  perhaps  little  less  common  in  foliage 
upon  very  short  internodes,  as  likewise  are  higher  ranked 
numbers ;  and  in  many  pine-cones  and  similar  structures  ^f 
and  f  £  phyllotaxy  may  be  readily  made  out.  This  actual  series, 
£,  £,  f ,  |,  &c.,  answers  to  and  may  be  expressed  by  the  con- 
tinued fraction,  %  +  \ 

t  +  i  +  |,  &c.2 


1  When  other  instances  are  detected,  they  are  found  to  belong  to  other 
series,  following  the  same  law,  such  as  the  rare  one  of  \,  \,  $ ,  T\. 

2  "  The  ultimate  values  of  these  continued  fractions  extended  infinitely 
are  complements  of  each  other,  as  their  successive  approximations  are,  and  are 
in  effect  the  same  fraction,  namely,  the  irrational  or  incommeasurate  inter- 
val which  is  supposed  to  be  the  perfect  form  of  the  spiral  arrangement. 
This  does,  in  fact,  possess  in  a  higher  degree  than  any  rational  fraction  the 
property  common  to  those  which  have  been  observed  hi  nature;  though 
practically,  or  so  far  as  observation  can  go,  this  higher  degree  is  a  mere 
refinement  of  theory.    For,  as  we  shall  find,  the  typical  irrational  inter- 
val differs  from  that  of  the  fraction  f  by  almost  exactly  \^6>  a  quantity 
much  less  than  can  be  observed  in  the  actual  angles  of  leaf -arrangements." 
"On  this  peculiar  arithmetical  property  ....  depends  the  geometrical  one, 
of  the  spiral  arrangement,  which  it  represents  ;  namely,  that  such  an  arrange- 
ment would  effect  the  most  thorough  and  rapid  distribution  of  the  leaves 
around  the  stem,  each  new  or  higher  leaf  falling  over  the  angular  space  be- 
tween the  two  older  ones  which  are  nearest  in  direction,  so  as  to  subdivide  it  in 
the  same  ratio  in  which  the  first  two,  or  any  two  successive  ones,  divide  the 
circumference.    But,  according  to  such  an  arrangement,  no  leaf  would  ever 
fall  exactly  over  any  other ;  and,  as  I  have  said,  we  have  no  evidence,  and 
could  have  none,  that  this  arrangement  actually  exists  in  nature.     To  realize 
simply  and  purely  the  property  of  the  most  thorough  distribution,  the  most 
complete  exposure  of  light  and  air  around  the  stem,  and  the  most  ample 
elbow-room,  or  space  for  expansion  in  the  bud,  is  to  realize  a  property  that 
exists  separately  only  in  abstraction,  like  a  line  without  breadth.   Neverthe- 
less, practically,  and  so  far  as  observation  can  go,  we  find  that  the  fractions 
|  and  -fa,  ^T,  &c.,  which  are  all  indistinguishable  as  measured  values  in  the 
plant,  do  actually  realize  this  property  with  all  needful  accuracy.     Thus, 
f  =  0.375,  •&  =  0.385,  and  &  ==  0.381,  and  differ  from  k  [the  ultimate  value 
to  which  the  fractions  of  this  series  approximate,  or  what  is  supposed  to  be 
the  type-form  of  them]  by  —0.007,  +0.003,  and  —0.001  respectively  ;  or  they 
all  differ  by  inappreciable  values  from  the  quantity  which  might  therefore  be 
made  to  stand  for  all  of  them.     But,  in  putting  k  for  all  the  values  of  the 
series  after  the  first  three,  it  should  be  with  the  understanding  that  it  is  not 
so  employed  in  its  capacity  as  the  grand  type,  or  source  of  the  distributive 
character  which  they  have,  —  in  its  capacity  as  an  irrational  fraction,  —  but 
simply  as  being  indistinguishable  practically  from  those  rational  ones."  — 
Chaucey  Wright,  in  Mem.  Amer.  Acad.  ix.  387-390. 


126  PHYLLOTAXY  OR  LEAF-ARRANGEMENT, 

241 ,  The  successive  grades  of  angular  divergence  of  alternate 
leaves,  as  expressed  in  degrees,  are 

i  =  180°  f  =  144°          Vk  =  138°  27'  41.54" 

|  =  120°  f  =  135°          £.  =  137°  8'  34.29" 

and  so  on  ;  and  beyond,  if  not  in  the  latter  cases,  the  differences 
become  quite  too  small  for  determination  by  inspection.  They 
all  fall  within  the  £  and  £  as  to  amount  of  divergence ;  and  they 
form  a  series  converging  to  a  deduced  typical  angle  of  137°  30' 
28",  which,  being  irrational  to  the  circumference,  would  place 
no  leaf  exactly  over  any  preceding  one,  but  alternately  and 
more  and  more  slightly  on  one  and  the  other  side  of  the  vertical, 
and  so  on,  in  an  endless  spiral.  That  is,  according  to  Bravais, 
the  ranks  in  the  higher  grades  tend  to  become  curviserial,  or 
actually  become  so;  while  in  the  lower  grades  they  are  obviously 
rectiserial.  Unless,  indeed,  there  is  some  torsion  of  the  axis, 
by  which  the  vertical  ranks  are  rendered  oblique,  as  is  often  the 
case  in  cones  of  the  Norway  Spruce.  But,  apart  from  this,  the 
difference  between  rectiserial  in  a  high  order  and  curviserial 
soon  becomes  inappreciable.  Any  and  all  of  the  higher  grades, 
and  practically  one  as  low  as  the  f ,  secures  the  utility  of  the 
theoretical  angle,  viz.,  that  "  by  which  the  leaves  would  be  dis- 
tributed most  thoroughly  and  rapidly  around  the  stem,  exposed 
most  completely  to  light  and  air,  and  provided  with  the  greatest 
freedom  for  symmetrical  expansion,  together  with  a  compact 
arrangement  in  the  bud."  Even  in  the  simpler  grades  of  com- 
monest occurrence,  each  leaf  (according  to  Wright)  is  so  placed 
over  the  space  between  older  leaves  nearest  in  direction  to  it  as 
always  to  fall  near  the  middle  of  the  space,  until  the  circuit  is 
completed,  when  the  new  leaf  is  placed  over  an  old  one.1 

242.  It  is  to  be  noted  that  the  distichous  or  £  variety  gives 
the  maximum  divergence,  viz.  180°,  and  that  the  tristichous  or 
£  gives  the  least,  or  120°  ;  that  of  the  pentastichous  or  £  is  nearly 
the  mean  between  the  first  two  ;  that  of  the  f ,  nearly  the  mean 
between  the  two  preceding,  &c.     The  disadvantage  of  the  two- 
ranked  arrangement  is  that  the  leaves  are  soon  superposed  and 
so  overshadow  each  other.     This  is  commonly  obviated  by  the 
length  of  the  internodes,  which  is  apt  to  be  much  greater  in  this 
than  in  the  more  complex  arrangements,  therefore  placing  them 
vertically  farther  apart ;  or  else,  as  in  Elms,  Beeches,  and  the 

1  This  corresponds  with  Hofmeister's  general  rule,  that  "new  lateral 
members  have  their  origin  above  the  widest  gaps  between  the  insertions  of 
the  nearest  older  members."  Yet  the  fact  that  the  character  of  the  leaf- 
arrangement  is  laid  down  at  the  beginning  in  the  bud  does  not  go  far  in  the 
way  of  the  mechanical  explanation  which  he  invokes- 


DISTRIBUTION    OF   LEAVES    ON   THE   STEM.  127 

like,  the  branchlets  take  a  horizontal  position  and  the  peti- 
oles a  quarter  twist,  which  gives  full  exposure  of  the  upper 
face  of  all  the  leaves  to  the  light.  The  £  and  f ,  with  dimin- 
ished divergence,  increase  the  number  of  ranks ;  the  f  and  all 
beyond,  with  mean  divergence  of  successive  leaves,  effect  a  more 
thorough  distribution,  but  with  less  and  less  angular  distance 
between  the  vertical  ranks. 

242°.  The  helix  or  primitive  spiral  upon  which  the  leaves 
successively  originate  ascends,  sometimes  from  left  to  right, 
sometimes  from  right  to  left,1  commonly  without  change  on  the 
same  axis,  and  prevailingly  uniform  in  the  same  species ;  but 
occasionally  both  directions  occur  in  the  same  individual.  The 
earliest  leaves  of  a  stem  or  branch,  or  the  last,  are  often  on  a 
different  order  from  the  rest ;  or  (as  already  stated)  the  spiral 
may  change  into  the  cyclical,  or  vice  versa. 

243.  The  relation  of  the  phyllotaxy  of  a  branch  to  the  leaf 
from  the  axil  of  which  the  branch  springs  is  somewhat  various. 
But  in  Dicotyledons,  the  first  leaf  or  the  first  pair  of  the  branch  is 
mostly  transverse  ;  that  is,  the  first  leaves  of  the  branch  stand  to 
the  right  or  left  of  the  subtending  leaf.     In  Monocotyledons,  the 
first  branch-leaf  is  usually  parallel  to  and  facing  the  subtending 
leaf,  as  shown  in  Fig.  304. 

244.  When  the  internodes  are  considerably  lengthened,  the 
normal  superposition  of  leaves  is  not  rarely  obscured  by  torsion 
of  the  axis  :  indeed,  this  may  equally  occur  in  short  internodes, 
sometimes  irregularly  or  in  opposite  directions,  sometimes  uni- 
formly in  one.     Thus,  in  Pandanus  utilis,  or  Screw-Pine,  of 
tristichous  arrangement,  the  three  compact  vertical  ranks  be- 
come strongly  spiral  by  a  continuous  torsion  of  the  axis.     The 
later  leaves  of  Baptisia  perfoliata,  which  are  normally  distichous, 
become  one-ranked  by  an  alternate  twist,  right  and  left,  of  the 
successive  internodes. 

245.  When  the  internodes  are  short,  so  that  the  leaves  approx- 
imate or  overlap,  it  is  difficult  or  impossible  to  trace  the  suc- 
cession of  the  leaves  on  the  primitive  spiral,  but  it  is  easy  to 
see  which  are  superposed.     The  particular  phyllotaxy  may  then 
be  determined  by  counting  the  vertical  ranks,  which  gives  the 
denominator  of  the   fraction.     But   in  compact  arrangements 
these  vertical  ranks  are  commonly  less  manifest  than  certain 
oblique  ranks,  which  are  seen  to  wind  round  the  axis  in  oppo- 
site directions.  (See  Fig.  245,  246.)     These  are  termed  second- 
ary spirals,  also   by  some  parastichies.     These   oblique   spiral 

1  That  is,  of  the  observer  and  as  seen  from  without.    See  p.  51,  foot-note. 


128  PHYLLOTAXY,  OB    LEAF-ARRANGEMENT. 

ranks  are  a  necessary  consequence  of  the  regular  ascending 
arrangement  of  parts  with  equal  intervals  over  the  circumference 
of  the  axis  ;  and,  if  the  leaves  are  numbered  consecutively,  their 
numbers  will  necessarily  stand  in  arithmetical  progression  on  the 
oblique  ranks,  and  have  certain  obvious  relations  with  the  pri- 
mary spiral  which  originates  them,  as  will  be  seen  by  projecting 
them  on  a  vertical  plane. 

245°.  Take,  for  example,  the  f  arrangement,  where,  as  in  the 
diagram  annexed  to  Fig.  244,  the  primitive  spiral,  written  on  a 
plane  surface,  appears  in  the  numbers,  1,  2,  3,  4,  5,  6,  and  so 
on  :  the  vertical  ranks  thus  formed  are  necessarily  the  numbers 
1-6-11  ;  4-9-14  ;  2-7-12  ;  5-10-15  ;  and  3-8-13.  But  two 
parallel  oblique  ranks  are  equally  apparent,  viz.  1-3-5,  which, 
if  we  coil  the  diagram,  will  be  continued  into  7-9-11-13-15  ;  and 
also  the  2-4-6-8-10  continues  into  12-14,  and  so  on,  if  the  axis 
be  prolonged.  Here  the  circumference  is  occupied  by  two  secon- 
dary left-hand  series,  and  we  notice  that  the  common  difference 
in  the  sequence  of  numbers  is  two ;  that  is,  the  number  of  the 
parallel  secondary  spirals  is  the  same  as  the  common  difference  of 
the  numbers  on  the  leaves  that  compose  them.  Again,  there  are 
other  parallel  secondary  spiral  ranks,  three  in  number,  which 
ascend  to  the  right ;  viz.  1-4-7,  continued  into  10-13  ;  3-6-9-12, 
continued  into  15 ;  and  5—8—11—14,  &c. ;  where  again  the  common 
difference,  3,  accords  with  the  number  of  such  ranks.  This  fixed 
relation  enables  us  to  lay  down  the  proper  numbers  on  the  leaves, 
when  they  are  too  crowded  for  directly  following  their  succes- 
sion, and  thus  to  ascertain  the  order  of  the  primary  spiral  series 
by  noticing  what  numbers  come  to  be  superposed  in  the  verti- 
cal ranks.  Thus,  in  the  small  cone  of  the  American  Larch 
(Fig.  244) ,  which  usually  completes  only  three  heights  of  leaves, 
the  lowest,  highest,  and  a  middle  one  make  a  vertical  row 
which  faces  the  observer.  Marking  this  first  scale  1 ,  and  count- 
ing the  parallel  secondary  spirals  that  wind  to  the  left,  we  find 
that  two  occupy  the  whole  circumference.  From  1,  we  number 
on  the  scales  of  that  spiral  3-5-7,  and  so  on,  adding  the  com- 
mon difference  2,  at  each  step.  Again,  counting  from  the  base 
the  right-hand  secondary  spirals,  we  find  three  of  them,  and 
therefore  proceed  to  number  the  lowest  one  by  adding  this  com- 
mon difference,  viz.  1-4-7-10 ;  then,  passing  to  the  next,  on 
which  the  No.  3  has  already  been  fixed,  we  carry  on  that  se- 
quence, 6-9,  &c. ;  and  on  the  third,  where  No.  5  is  already 
fixed,  we  continue  the  numbering,  8-11,  &c.  This  gives  us  in 
the  vertical  rank  to  which  No.  1  belongs  the  sequence  1-6-1 1 , 
showing  that  the  phyllotaxy  is  of  the  five-ranked,  or  f  order. 


DISTRIBUTION    OF   LEAVES    ON    THE   STEM.  129 

It  is  further  noticeable  that  the  smaller  number  of  parallel  sec- 
ondary spirals,  2,  agrees  with  the  numerator  of  the  fraction  in 
this  the  f  arrangement ;  and  that  this  number,  added  to  that  of 
the  parallel  secondary  spirals  which  wind  in  the  opposite  direction, 
viz.  3,  gives  the  denominator  of  the  fraction.  This  holds  good 
throughout ;  so  that  we  have  only  to  count  the  number  of  par- 
allel secondary  spirals  in  the  two  directions,  and  assume  the 
smaller  number  as  the  numerator,  and  the  sum  of  this  and  the 
larger  number  as  the  denominator,  of  the  fraction  which  ex- 
presses the  angular  divergence  sought.  For  this,  we  must,  how- 
ever, take  the  order  of  secondary  spirals  nearest  the  vertical 
rank  in  each  direction,  when  there  are  more  than  two,  as  in  all 
the  higher  forms.  But,  in  all,  it  is  necessary  to  count  only  the 
most  manifest  seconda^  spiral  of  each  direction  in  order  to 
lay  down  the  proper  number  on  the  leaves  or  scales,  and  so  deter- 
mine the  phyllotaxy.1  In  a  rosette  of  the  leaves  of  Houseleek 
(Fig.  245)  and  a  cone  of  Pinus  Strobus  (Fig.  246),  the  num- 
bers which  can  be  seen  at  one  view  are  appended,  and  in  the 
latter  the  conspicuous  secondan-  spirals  are  indicated  :  one  to 
left  with  a  common  difference  of  5  ;  and  two  to  the  right,  of 
which  the  most  depressed  and  prominent  has  the  common  dif- 
ference of  3,  the  other,  nearest  the  vertical,  the  common  differ- 
ence 8.  The  14th  leaf  is  superposed  to  the  first,  indicating  the 
•fe  arrangement.  The  same  conclusion  is  derived  from  the  num- 
ber of  the  higher  spirals,  the  smaller  5  for  the  numerator,  and 
this  added  to  8  for  the  denominator.  The  mathematical  discus- 
sion of  these  relations,  and  of  the  whole  subject  of  phyllotaxy, 
leads  into  interesting  fields.  But  this  sketch  may  suffice  for 
botanical  uses. 

246.  Relations  of  Whorls  to  Spirals.  Verticillate  and  alternate 
phyllotaxy,  or  whorls  and  spirals,  in  all  complete  exemplifica- 
tions, are  to  be  considered  morphologicall}-  as  distinct  modes, 
not  to  be  practically  homologized  into  one.  Nevertheless,  transi- 
tions between  the  two,  and  abrupt  changes  from  one  to  the  other 
on  the  same  axis  are  not  uncommon,  the  former  especially  in 
the  foliage,  the  latter  in  the  blossom.  If  the  spiral  be  assumed 
as  the  fundamental  oi'der,  it  is  not  difficult  to  form  a  clear  con- 
ception as  to  how  such  changes  come  to  pass.  A  single  whorl 

1  In  applying  this  method  to  the  determination  of  the  phyllotaxy  of  a 
cone,  or  any  such  assemblage  of  leaves,  the  student  should  be  warned  that, 
although  the  cones  of  Pines  and  Firs  are  all  normally  on  the  alternate  plan 
(while  those  of  Cypresses  are  on  the  verticillate),  yet  in  individual  cases 
(common  in  Norway  Spruce)  the  cone  is  plainly  ma<V  up  of  pairs  of  oppo- 
site scales  which  are  spirally  arranged.  See  note  under  236. 


130  PHYLLOTAXY,   OK  LEAF-ARRANGEMENT. 

may  most  naturally  be  produced  by  the  non-development  of  the 
internodes  between  any  two,  three,  or  more  alternate  leaves. 
Two  proximate  distichously  alternate  leaves  would  thus  form  a 
pair ;  the  three  leaves  belonging  to  one  turn  of  the  spiral  in  the 
tristichous  (^)  arrangement  would  compose  a  trimerous  whorl ; 
the  five  leaves  of  the  two  turns  in  the  pentastichous  (f )  arrange- 
ment, a  5-merous  whorl,  &c.  Verifications  of  this  conception,  by 
whorls  breaking  up  or  reverting  to  spirals,  are  occasionally  met 
with,  and  the  successive  overlapping  in  spiral  order  of  the 
members  of  a  trimerous  or  pentamerous  whorl  is  very  common. 
The  few  instances  among  phaenogamous  plants  in  which  the 
leaves  are  opposite  and  all  in  the  same  plane  1  (that  is,  the  suc- 
cessive pairs  superposed)  may  be  deduced  from  the  distichous 
alternate  mode  becoming  opposite  without  further  change,  by 
the  simple  suppression  of  alternate  internodes.  The  frequent 
disjunction  of  the  members  of  the  pair  in  similar  and  analogous 
cases  goes  to  confirm  this  view.  But  the  characteristic  of  whorls 
ordinarily  is  that  proximate  whorls  alternate,  that  pairs  de- 
cussate. We  cannot  homologize  this  with  spiral  phyllotaxy ; 
for  in  this  lies  the  fundamental  difference  between  the  two  plans. 
We  can  explain  it  only  by  a  reference  to  Hofmeister's  law,  which 
generally  governs  leaf-origination  as  to  position,  namely,  that 
succeeding  leaves  appear  directly  above  the  intervals  between 
the  nearest  preceding  (241,  note)  :  this  gives  decussation  or 
alternation  of  successive  pairs  or  whorls.2 

247.  Hypothesis  of  the  origin  of  both.  Instead  of  regarding 
the  spiral  path  on  the  stem  which  connects  successive  alternate 
leaves  as  a  purely  formal  representation,  it  may  be  conceived  to 
be  the  line  along  which  the  members  in  some  original  form  were 
physically  connected,  in  the  manner  of  a  leaf-like  expansion 

1  As  in  Loranthus  Europaeus,  &c.,  according  to  Braun.     See  236,  note. 

2  This  renders  the  verticillate  an  advantageous  arrangement,  perhaps  no 
less  so  than  the  distribution  which  spiral  phyllotaxy  effects.     Both  must  be 
considered  to  have  been  determined  by  and  for  their  respective  utilities,  and 
to  have  been  independent  determinations.     For  "  there  is  no  continuity  or 
principle  of  connection  between  spiral  arrangements  and  whorls  "  (Chauncey 
Wright) ;  since,  although  individual  whorls  are  easily  reducible  to  spirals, 
each  succession  is  an  absolute  break  of  that  system. 

As  whorls  of  four  members  often  (as  especially  in  calyx,  bracts,  &c.)  may 
and  sometimes  should  be  viewed  as  two  approximate  pairs,  so  even  the  spiral 
of  five  members,  as  in  a  quincuncial  calyx,  has  been  conceived  to  consist  of 
two  whorls,  one  of  two,  the  other  of  three  leaves,  the  second  alternating  with 
the  first  as  nearly  as  possible.  But  this  appears  far-fetched  and  of  loose 
application.  It  is  much  clearer  as  well  as  simpler  to  regard  the  alternate 
as  the  fundamental  phyllotaxy,  and  to  deduce  individual  whorls  from  spirals, 
if  need  be,  rather  than  to  imagine  spirals  as  somehow  evolved  from  whorls. 


DISTEIBUTION  OF   LEAVES   ON   THE   STEM.  131 

resembling  a  spiral  stairway.  Upon  this  supposition,  the  leaves 
would  be  the  relics,  or  rather  the  advantageous  results,  of  the 
segmentation  of  such  a  frond-like 
expansion,  the  segments  separated 
through  the  development  of  the 
stem  in  length  and  firmness,  and 
modified  in  the  various  adaptations 
to  the  conditions  of  higher  vege- 
table life ;  even  as  leaves  themselves 
are  modified  into  tendrils,  bud- 
scales,  petals,  or  other  usefully  specialized  structures.  The  type 
on  this  conception  would  be  a  frond,  consisting  of 
an  elongating  axis  with  a  continuous  leaf-blade  on 
one  side,  and  this  taking  a  spirally  twisted  form. 
But  the  frond  of  Fucaceous  Algae,  Hepaticae,  and  the 
like,  is  two-bladed.  While  a  one-bladed  frond,  or 
with  one  blade  suppressed,  might  be  the  original  of 
alternate-leaved  spirals,  the  two-bladed  frond,  simi- 
larly broken  up,  would  give  rise  to  the  opposite  or 
other  varieties  of  verticillate  arrangement.1 

248.  Fascicled  Leaves  need  to  be  mentioned  here, 
in  order  that  they  may  be  excluded  from  phyllotaxy. 
They  are  simply  a  cluster  or  tuft  of  leaves,  belonging 
to  more  than  one  node,  and  left  in  a  crowded  con- 
dition because  the  internodes  do  not  lengthen.  They 
may  belong  either  to  the  alternate  or  the  verticillate 
series.  In  Barberry  and  in  the  Larch  (Fig.  247), 
they  are  evidently  alternate  ;  and  they  may  be  inferred 
to  be  so  in  Pines  (Fig.  248) ,  or  even  may  be  seen  to 
be  so  in  the  bud-scales  which  form  the  sheath  sur- 
rounding the  base  of  the  2,  3,  or  5  foliage-leaves. 
In  Junipers,  the  leaves  of  the  fascicles  are  in  the 
verticillate  order. 

1  This  is  the  conception  of  the  late  Chauncey  Wright.  See  his  elaborate 
and  most  suggestive  essay  in  Mem.  Amer.  Acad.  Arts  and  Sciences,  ser.  2, 
ix.  379,  mainly  reprinted  in  Philosophical  Discussions  (posthumous),  296-328, 
in  which  the  whole  subject  of  phyllotaxy  is  acutely  discussed,  especially 
in  its  relation  to  questions  of  origin  and  developed  utilities.  His  conception 

FIG.  247.  Piece  of  a  branch  of  the  Larch,  with  two  fascicles  of  leaves,  i.  e.  two  very 
short  and  stout  branchlets,  bearing  scars  of  former  leaves  or  bud-scales  below,  and  a 
dense  cluster  of  leaves  of  the  season  at  summit.  The  main  axis  bears  scars  from  which 
the  alternate  leaves  of  the  developed  axis  of  the  preceding  year  separated. 

FI&.  248.  Piece  of  a  branch  of  Pitch  Pine,  with  three  leaves  in  a  fascicle  or  bundle 
in  the  axil  of  a  thin  scale  (a)  which  answers  to  a  leaf  of  the  main  axis.  The  bundle  is 
surrounded  at  thu  base  by  a  short  sheath,  formed  of  the  delicate  scales  of  the  axillary 
bud,  of  which  the  three  leaves  are  the  developed  foliage. 


132  PHYLLOTAXY,  OR   LEAF-ARRANGEMENT. 


SECTION  II.     DISPOSITION  OF  LEAVES  IN  THE  BUD. 

249.  Vernation  and  Estivation  are  terms  in  general  use,  under 
which  the  disposition  of  leaves  in  the  bud  is  treated.  The  first 
relates  to  ordinary  leaves  in  this  early  condition  ;  the  second,  to 
the  parts  of  a  flower-bud ;  not,  however,  as  respects  insertion, 
or  position  on  the  axis,  which  is  phyllotaxy  (231),  but  as  to 

the  wa}-s  in  which  they 
are  coiled,  folded,  over- 
lapped, &c.,  either  joer  se 
or  inter  se.  Prcefoliation 
and  Prcefloration  are 
etymologically  better 
terms,  substituted  by 
Richard.1 

250.  The  descriptive 
terms  which  relate  to 
indiAddual  leaves  or 
parts,  whether  of  foli- 
age or  blossom,  mostly  range  themselves  under  the  heads  of 
plications  or  of  enrolling,  and  are  such  as  the  following,  the 
sectional  diagrams  of  which  are  copied  from  the  original  figures 

would  make  the  two  plans  equally  primordial.  But  the  freedom  with 
which  these  actually  interchange  on  the  same  axis  greatly  favors  the  less 
hypothetical  view  that  whorls  may  be  condensed  spirals.  This  assumes 
only  the  well-known  fact  that  internodes  may  be  completely  non-developed. 

1  Better  formed  and  more  expressive  terms:  but  the  Linnasan  ones  are 
most  in  use,  and,  though  fanciful,  are  not  misleading.  In  English  description, 
it  is  as  convenient  and  equally  terse  to  say  that  the  parts  are  imbricate,  val- 
vate,  &c.,  "  in  the  bud."  Linnaeus,  in  the  Philosophia  Botanica,  described 
these  dispositions  of  leaves  in  the  bud  under  the  term  Foliatio,  — not  a  happy 
name,  —  but  did  not  treat  of  them  in  the  flower-bud.  Later,  in  Termini 
Botanici  (Amoen.  Acad.  vi.  1762,  reprinted  by  Giseke  in  1781),  he  intro- 
duced the  words  Vernatio  and  ^Estivatio  in  their  now  current  botanical  sense, 
to  designate,  not  the  time  of  leafing  and  of  flowering  (spring  and  summer 
condition),  but  the  disposition  of  the  parts  in  the  leaf-bud  and  flower-bud 
(at  least  of  the  petals)  as  respects  foldings,  coiling,  &c.,  of  single  parts,  and 
modes  of  overlapping  or  otherwise  of  contiguous  parts.  The  terminology 
as  regards  single  leaves,  Linnaeus  fixed  nearly  as  it  now  remains.  That  of 
leaves  or  their  homologues  in  connection,  and  as  respects  the  flower-bud,  was 
very  imperfectly  developed  until  its  importance  (and  much  of  its  termi- 
nology) was  indicated  by  Robert  Brown,  in  his  memoir  on  Proteaceae,  1809, 
in  the  Prodromus  a  year  later,  and  in  other  publications. 

Ptyxis  (the  Greek  name)  is  coming  into  use  as  a  general  term  for  the 
folding,  &c.,  of  single  parts. 

FIG.  249-254.  I.innaean  diagrams  of  sections  of  leaves  in  the  bud  249.  Conduph- 
cate.  250.  Plicate  or  plaited  251.  Convolute.  252.  Revolute.  253.  Involute.  254. 
Circinate  or  Circiiial 


VERNATION,    OR    PR^EFOLIATION.  133 

in  the  Philosophia  Botanica  of  Linnaeus.  They  were  applied 
onty  to  foliage,  but  they  are  equally  applicable  to  floral  parts. 
Leaves,  and  all  homologous  or  similar  organs,  if  not  simply 
plane,  will  be  either  bent  or  folded  or  else  more  or  less  rolled  up 
in  the  bud.  The  first  three  of  the  following  terms  relate  to  the 
former,  the  remaining  terms  to  the  latter.  They  are  as  to  the 
mode  of  packing 

Plicate  or  Plaited  (Fig.  250) ,  when  folded  on  the  several  ribs, 
in  the  manner  of  a  closed  fan,  as  in  Maple  and  Currant.  This 
occurs  only  in  certain  palmately  veined  or  nerved  leaves. 

Conduplicate  (Fig.  249),  when  folded  lengthwise,  or  doubled 
up  flat  on  the  midrib,  as  in  Magnolia ;  a  verj-  common  mode. 
The  upper  face  of  the  leaf  is  always  within. 

Reclinate  or  Inflexed,  when  the  upper  part  is  bent  on  the 
lower,  or  the  blade  on  the  petiole,  as  in  the  Tulip-tree  (the  blade 
of  which  is  also  conduplicate) . 

Convolute  (Fig.  251),  when  rolled  up  from  one  margin,  i.e. 
one  margin  within  the  coil,  the  other  without,  as  in  Apricot 
and  Cherry. 

Involute  (Fig.  253) ,  both  margins  rolled  toward  the  midrib 
on  the  upper  face,  as  the  leaves  of  Water  Lily,  Violet,  &c.  ; 
also  the  petals  of  Steironema  and  Tremandra. 

Revolute  (Fig.  252),  similarly  rolled  backwai'd  from  both 
margins,  as  the  leaves  of  Azalea  and  Rosemary. 

Gircinal  or  Circinate  (Fig.  254) ,  when  coiled  from  the  apex 
downward,  as  the  leaves  of  Drosera  and  the  fronds  of  all  the 
true  Ferns. 

Corrugate  or  Crumpled,  as  the  petals  of  a  Poppy,  applies 
to  the  irregular  crumpling  of  the  otherwise  plane  corolla-leaves. 
This  is  a  consequence  of  rapid  growth  in  length  and  breadth 
in  a  confined  space. 

251.  The  Ptyxis  (or  folding,  &c.)  of  an  individual  leaf,  of 
which  the  foregoing  modifications  are  the  principal,  should  be 
distinguished  from  the  arrangement  in  the  bud  of  the  leaves  of 
a  circle  or  spiral  in  respect  to  each  other.     The  interest  of  the 
latter  centres  in  the  flower-bud,  i.  e.  in  aestivation.     To  this  the 
following  exposition  is  devoted,  although  sometimes  applicable 
to  leaf-buds  also.1 

252.  The  disposition   of  parts  in   aestivation,  in  respect  to 
each  other,  is  the  result  partly  of  their  relative  insertion,  that  is 

1  In  the  succeeding  paragraphs,  it  becomes  necessary  to  presuppose  so 
much  knowledge  of  the  flower  as  is  implied  in  the  free  use  of  such  terms  as 
calyx  and  corolla,  sepals  or  calyx-leaves,  and  petals  or  corolla-leaves.  See, 
if  need  be,  Chapter  VI.  Sect.  I. 


184  PHYLLOTAXY,  OR  LEAF-ARRANGEMENT. 

their  phyllotaxy,  and  partly  of  the  way  in  which  they  comport 
when  their  margins  meet  in  growth.  Those  leaves  which  are 
within,  or  of  higher  insertion  on  the  axis,  will  almost  necessarily 
be  enclosed  or  overlapped :  those  which  are  members  strictly 
of  the  same  whorl  or  cycle  may  fail  to  come  into  contact,  or 
may  meet  without  overlapping  at  the  contiguous  margins  or 
apex ;  yet  they  may  be  overlapped,  since  they  may  have  grown 
unequally  or  some  a  little  earlier  than  their  fellows.  Conse- 
quently, no  perfectly  clear  line  can  be  drawn  in  the  flower  between 
cycles  and  spirals  except  by  their  mode  of  succession.  More- 
over, aestivation  strictly  so  called  should  be  concerned  only 
with  the  disposition  among  themselves  of  the  several  members 
of  one  whorl,  or  of  one  complete  spiral.  So  the  alternation  of 
contiguous  whorls,  as  of  the  three  inner  with  the  three  outer 
flower-leaves  of  a  Lily  or  a  Tulip  (the  alternative  aestivation  of 
DeCandolle) ,  is  a  matter  of  phyllotaxy,  not  of  aestivation.  The 
latter  is  properly  concerned  only  with  the  relations  of  each  three 
leaves  to  each  other.1 

253.  The  proper  aestivations  may  be  classified  into  those  in 
which  the  parts  do  not  overlap,  and  those  in  which  they  do.     Of 

the  first,  there  are  two  kinds,  the  open  (cest. 

aperta)  and  the  valvate,  both  characterized  and 

named  by  Brown.2    Of  the  second,  there  is 
i^]|  \\  one  leading  kind,  the  imbricate  (adopted  by 
'  j)  Brown  from  Linnaeus) ,  with  subordinate  modi- 
<v  ^»<^       fications.8      Accordingly,   the   aestivation   is 

said  to  be 

255  254.    Open  or   Indeterminate  (cest.  aperta), 

when  the  parts  do  not  come  into  contact  in  the  bud,  so  as  to 


1  The  same  applies  to  the  two  sets  of  sepals  and  of  petals  in  Barberry,  in 
Menispermum,  and  of  the  petals  in  Poppy,  &c.  (359). 

,  2  Linnaeus,  indeed,  has,  "  ^Estivatio  valvata,  si  petala  se  expansura  instar 
glumae  graminis  ponuntur,"  —  the  name,  but  not  the  thing:  the  glumes  of 
grasses  are  not  valvate  in  the  botanical  sense.  So  the  term  as  to  its  proper 
use  may  be  said  to  originate  with  R.  Brown. 

8  For  a  brief  discussion  of  "  ^Estivation  and  its  Terminology,"  see  Amer. 
Jour.  Sci.  ser.  3,  x.  339,  1875. 

As  to  names,  it  is  perhaps  more  correct  to  say  of  the  (estivation  that  it  is 
imbricative,  convolutive,  valvular,  &c.  (cest.  imbricativa,  convolutiva,  valvaris,  &c.), 
but  of  the  leaves  or  pieces,  that  they  are  imbricate,  convolute,  valvate,  &c.,  in 
aestivation  ;  but  such  precision  of  form  will  seldom  be  attended  to  in  botan- 
ical descriptions. 

FIG.  255.  Diagrammatic  cross  section  of  an  unopened  flower  of  Linden :  its  outer 
circle  of  floral  leaves  (sepals)  valvate  In  the  bud;  the  Inner  (petals)  between  convolute 
and  imbricate. 


AESTIVATION,   OR  PR^FLOEATION.  135 

cover  those  within.     The  most  familiar  case  is  that  of  the  petals 
of  Mignonette  and  the  whole  genus  Reseda. 

255.  Valvate  or  Valvular,  when  the  margins  meet  squarely  in  the 
oud,  without  any  overlapping,  like  the  valves  of  a  dehiscent  cap- 
sule.   Familiar  examples  are  afforded  by  the  calyx  of  the  Linden 
(Fig.  255)  ;  also  that  of  the  Mallow,  Rhamnus,  Fuchsia,  and  the 
whole  of  the  several  natural  orders  to  which  these  belong.     A 
modification  of  this,  caused  by  some  induplication  or  involution 
of  the  edges  of  the  individual  leaves,  occurs  in  most  species  of 
Clematis :  in  Clematis  Virginiana,  they 

merely  project   within    (yalvate-indupli- 

cate)  ;  in  Clematis   Viticella,    they   are 

conspicuously  involute  (valvate-involute) , 

or  valvate  with  margins  involute.    Some-          256 

tunes  (as  in  the  calyx  of  certain  Malvaceae)   the  joined  edges 

project  outwardly  (or  are  valvate  with  reduplicate  margins) ,  but 

only  slightly  so. 

256.  Imbricate  or  Tmbricative  is  the  general  name  for  aestiva^ 
tion  (or  vernation)  with  overlapping.     The  name  is  taken  from 
the  overlapping  of  tiles  or  shingles  on  a  roof,  so  as  to  break 
joints  or  cover  edges.     It  was  first  applied,  by  Linnaeus,   to 
leaves  or  scales  on  a  stem,  when  thickly  set  and  incumbent  in  suo- 
cessive  ranks  or  heights,  the  upper  partly  covered  by  those  next 
below.     The  involucre  of  an  Aster  or  of  the  common  Sunflower 
is  a  typical  illustration  ;  as  also  the  leaves  of  a  Camellia-flower, 
the  sepals  as  well  as  the  petals  ;  and  the  sepals  or  outer  leaves 


238 

of  a  Flax  or  a  Geranium-flower  afford  a  simpler  but  similar 
instance,  although,  from  the  parts  being  nearly  of  the  same  size 
and  at  the  same  height,  the  overlapping  is  lateral  instead  of 
obviously  from  below.  Fig.  258,  259,  and  the  outer  part  of 
260,  also  the  inner  leafy  circle  of  255,  illustrate  in  diagram  this 
true  and  simple  imbricative  aestivation  of  a  definite  number  of 

FIG.  256.  Valvate-imluplicate  flower-leaves  (calyx)  of  Clematis  Virginiana,  &c. 
257.  Valvate-involute,  as  in  C.  Viticella. 

FIG.  258-260.  Imbricate  {estivation :  258,  in  two  whorls  of  three  leaves  each  (calyx 
and  corolla);  260,  same  of  five  leaves  in  the  outer  circle,  those  of  the  inner  circle  con- 
volute; 259,  a  single  set  of  three  imbricated  leaves  (in  the  corolla  of  Magnolia), 
almost  completely  encircling  each  otber. 


136  PHYLLOTAXY,  OR   LEAF-ARRANGEMENT. 

parts.1  It  is  characteristic  of  it  that  some  parts  (one  or  more) 
are  wholly  exterior  or  covering  in  the  bud,  and  others  (one  at 
least)  interior  or  covered,  at  least  the  margins.  Imbricative 
aestivation,  it  will  be  seen,  naturally  attends  alternate  or  spiral 
phyllotaxy  (248,  and  see  Fig.  242,  243)  ;  and  if  it  be  main- 
tained that  these  sets  of  three,  five,  &c.,  in  the  blossom  are  not 
depressed  spirals,  but  whorls  or  cycles  (as  may  commonly  be 
the  case  in  the  corolla,  but  hardly  in  the  calyx) ,  it  is  not  less 
true  that  the  parts  are  apt  to  comport  themselves  in  the  exact 
manner  of  a  depressed  spiral.  The  kinds  of  regular  imbrication 
of  alternate  leaves,  &c.,  may  be  specified  by  the  terms  or  frac- 
tions expressive  of  the  particular  grade  of  phyllotaxy  (£,  ^,  £ , 
|,  &c.).  But  some  of  them  have  received  special  names,  which 
may  be  employed,  as  subordinate  to  the  general  denomination  of 
imbricate.  The  most  important  of  these  are  the 

Equitant,  where  leaves  override,  the  older  successively  astride 
the  next  younger.  The  typical  instance  is  that  of  ancipital  or 
two-ranked  (^)  conduplicate  leaves,  successively  clasping,  at  least 
next  the  base,  as  in  Iris,  Fig.  217.  In  what  Linnaeus  termed 
equitant-triquetrous  (well  seen  in  Fig.  240,  241),  the  leaves  are 
three-ranked  (being  of  the  £  order),  and  each  imperfectly 
conduplicate. 

Quincuncial  aestivation  (as  in  the  outer  part  of  Fig.  260)  is 
simply  the  imbricate  aestivation  of  five  leaves  (§),  in  which 
necessarily  the  first  and  second  are  external,  the  fourth  and  fifth 
internal,  and  the  third  with  one  margin  external,  where  it  over- 
lies the  fifth,  and  the  other  internal,  where  it  is  covered  by 
the  first. 

Alternative  aestivation,  as  already  stated  (252),  comes  from 
verticillate  or  cyclic  phyllotaxy,  and  the  alternation  of  successive 
whorls.  When  two  such  whorls,  say  of  three  leaves  each  (as  in 
Fig.  258) ,  are  so  condensed  or  combined  as  to  form  apparently 
one  set  or  circle  of  six  members  (as  in  the  flower-leaves  of  most 
Liliaceae) ,  three  members  alternate  with  and  are  covered  by  the 
other  three,  and  this  sort  of  imbricate  aestivation  is  produced. 
More  properly,  the  two  series  are  to  be  considered  separately. 
Where  the  parts  are  four  (as  in  Fig.  395) ,  the  normal  imbrica- 
tion is  decussate,  two  exterior  and  two  interior.  This  is  some- 

1  All  the  examples  referred  to  result  from  alternate  or  spiral  phyllotaxy, 
the  former  of  higher  series,  the  latter  of  the  £  (Fig.  258,  259),  and  of  the  $ 
(Fig.  260)  order.  Instead  of  separating  (with  DeCandolle  and  others)  the 
\  arrangement  as  different  in  kind  from  the  imbricate  (under  the  name  of 
quincuncial  aestivation),  we  should  count  it  as  a  typical  case.  Otherwise  the 
i  arrangement  might  equally  claim  a  generic  distinction,  also  the  f,  &c. 


AESTIVATION,   OR   PR^EFLORATION.  137 

times  a  clear  case  of  binary  instead  of  quaternary,  t. e.  to  be 
counted  as  two  pairs  of  opposite  leaves  ;  yet  it  may  be  a  single 
whorl  of  four,  notwithstanding  the  imbrication.  Or  these  four 
leaves  may  even,  in  some  cases,  be  regarded  as  a  portion  of  a 
depressed  spiral,  say  of  the  f  order  with  one  piece  omitted,  and 
the  others  adjusted  so  as  to  fill  the  space. 

257.  There  are  various  deviations  from  normally  imbricative 
aestivation,  especially  where  the  members  are  five,  occurring 
some  in  regular  but  more  in  irregular  flowers,  which  need  not  be 
referred  to  here.  One,  for  which  no  specific  name  is  requisite, 
is  a  case  merely  of  excessive  overlapping  in  the  regular  way ; 
namely,  where  each  piece  completely  and  concentrically  encloses 
the  next  interior,  as  shown  in  Fig.  259,  representing  three  petals 
of  Magnolia  Umbrella.  This  the  French  botanists  have  called 
convolute  aestivation,  because  the  individual  leaves  are  involute 
in  a  manner  approaching  the  convolute  vernation  of  Linnaeus. 
Another  is  the  Vexillar,  as  in  the  Pea  tribe  (Fig.  306),  where 
members  which  should  be  external  have  somehow  developed  as 
internal,  both  in  calyx  and  corolla.  A  third  (which  has  received 
the  usually  quite  meaningless  name  of  Cochlear,  spoon-like, 
and  is  also  that  to  which  most  French  botanists  singularly  re- 
strict the  name  of  imbricative)  is  a  state  exactly  intermediate 
between  the  quincuncially  imbricate  and  the  convolute  or 
©  •  © 


contorted.  In  it,  one  leaf  is  wholly  outside,  one  wholly  inside, 
and  three  with  one  margin  inside  and  the  other  outside.  It  occurs 
under  two  modifications,  viz.  with  the  innermost  leaf  remote  from 
the  outmost  (Fig.  261),  and  with  it  next  to  the  outermost  as  in 
Fig.  262.  In  view  of  the  intermediate  character,  we  had 
applied  to  this  the  somewhat  awkward  name  of  Convolute-imbri- 
cated To  bring  Fig.  261  back  to  the  quincuncially  imbricate 

1  It  would  not  be  amiss,  therefore,  to  name  one  of  these  modes,  viz.  that 
of  Fig.  261,  Subimbricate,  and  the  other,  Fig.  262,  Subconvolute.  George 

FIG.  261.  Quincuncial  imbricate  modified  toward  convolute  by  one  edge  of  the 
second  leaf  developing  inside  instead  of  outside  of  the  adjacent  edge  of  the  fourth. 

PIG.  262.  Convolute  modified  toward  imbricate  by  one  leaf  having  a  margin  inside 
instead  of  outside  its  neighbor. 

FIG.  263.  Convolute,  or  convolutive,  or  contorted  (twisted)  aestivation,  in  diagram. 
In  these  throe  diagrams,  the  dark  circle  above  represents  the  position  of  the  axis,  tho 
flowers  being  axillary. 


138  PHYLLOTAXY,   OK   LEAF-AKKANGEMENT. 

form,  we  have  only  to  reverse  a  single  overlapping  on  the  left- 
hand  side  of  the  figure.  To  restore  Fig.  262  to  the  convolute, 
we  have  only  to  reverse  a  single  overlapping  at  the  lower  right- 
hand  side.  Changes  like  these,  or  the  reverse,  are  not  rare  in 
several  species,  particularly  in  the  corolla.  The  normal  form 
and  the  deviation  often  occur  in  different  flowers  on  the  same 
individual,  thus  indicating  an  easy  passage  between  the  imbricate 
aestivation  in  the  proper  sense  and  the 

258.  Convolute,  otherwise  called  Obvolute  or  Contorted,  or 
Twisted,  Fig.  263,  and  inner  circle  of  Fig.  260.  Here  each  leaf 
successively  overlaps  a  preceding  and  is  overlapped  by  a  following 
one,  all  having  a  slight  and  equal  obliquity  of  position,  so  that 
all  alike  have  an  exterior  and  an  interior  (or  a  covering  and  a 
covered)  margin,  and  all  appear  to  be  as  it  were  rolled  up  to- 
gether. This  is  strikingl}'  so  when  the  parts  are 
broad  and  much  overlapped,  as  in  Fig.  264.  Brown 
included  this  among  the  forms  of  imbricate  aestiva- 
tion, and  so  does  Eichler,  particularly  distinguishing 
it,  however,  under  the  name  here  preferred.  The 
264  occasional  transitions  would  justify  such  classifica- 

tion. But  in  most  cases  it  is  so  uniform,  and  in  the  corolla  so 
completely  characteristic  of  whole  families  (such  as  Malvaceae, 
Onagraceae,  Apocynacese,  Gentianaceae,  Polemoniaceae,  &c.),  and 
is  so  distinct  in  its  nature,  that  it  may  well  take  rank  among 
the  primary  kinds  of  aestivation.  As  to  its  nature,  it  is  evident 
that  while  the  imbricate  mode  (at  least  the  ternary,  quinary, 
&c.)  indicates  or  imitates  spiral  phyllotaxy  (some  members  be- 
ing within  or  with  higher  insertion  than  others) ,  the  convolute 
and  the  valvate  (having  all  the  members  of  the  series  on  the 
same  plane)  answer  to  verticillate  phyllotaxy,  or  to  whorls 
instead  of  depressed  spirals.1  The  name  which  this  mode  of 


Henslow,  in  Trans.  Linn.  Soc.  ser.  2,  i.  178,  proposes  to  call  the  former  half- 
imbricate;  the  latter  (following  the  faulty  French  example)  is  his  imbricate 
proper. 

The  subimbricate  mode  has  two  varieties,  distinguished  by  Eichler  (in 
Bliithendiagramme)  as  ascensive,  when  the  lower  or  anterior  (t.  e.  the  pieces 
next  the  subtending  bract  or  leaf)  are  successively  exterior  (as  in  Fig.  261), 
and  descensive,  when  the  covering  is  from  the  upper  side,  t.  e.  from  the  side 
next  the  axis. 

1  Still,  as  those  members  of  a  quincunx  which  normally  should  be  wholly 
external  do  sometimes  become  internal  during  their  development  in  the  bud, 
similar  changes  may  be  conceived  to  change  a  quincuncial  into  a  convolute 
disposition  ;  but,  to  effect  this,  three  out  of  the  five  overlappings  would  have 
to  be  reversed. 

FIG.  264.    Convolute  (also  called  contorted)  aestivation  of  a  corolla. 


AESTIVATION,   OR   PR^EFLORATION.  139 

estivation  ought  to  bear  is  not  yet  well  settled,  but  that  of  con- 
volute, here  preferred,  will  probably  prevail.1 

259.  In  recapitulation,  these  principal  forms  of  the  aestivation 
of  floral  circles  may  be  classified  in  a  synopsis.     They  are :  I. 
those  not  closed  =  open  or  indeterminate :  II.  closed  ;  and  these 
1,  with  the  margins  not  overlapping  =  valvate  ;  2,  with  margins 
overlapping ;  a,  one  or  more  with  both  margins  covered  =  imbri- 
cate ;  5,  all  with  one  margin  covered,  the  other  uncovered  = 
convolute. 

260.  Plicate  or  Plaited,  when  applied  to  the  flower-bud  as  a 
whole,  is  in  a  somewhat  different  category.     The  term  is  here 
used  for  the  plaiting 

of  a  tube  or  cup, 
composed  of  a  circle 
of  leaves  combined 
into  one  body.  It 

is   well   marked   in  265  266 

the  corolla  of  Convolvulus  and  of  Datura,  and 
in  most  of  the  order  to  which  these  belong.  In 
Campanula,  these  plaits  are  all  outwardly  sa- 
lient and  straight  (Fig.  265)  ;  in  the  corolla  of 
most  Gentians,  the  plaits  are  internal  and  straight. 
In  Convolvulus  and  Datura  (Fig.  266-268), 
the  narrow  plaits  overlap  one  another  in  a  con- 
volute way,  when  they  are  said  to  be  Supervolute. 
In  the  common  Morning  Glory  and  some  other 
species  of  Ipomoea,  these  plaits  are  besides  spirally  twisted  or 

1  See  article  entitled  "  -(Estivation  and  its  Terminology,"  above  referred  to. 
The  earliest  name  is  Obvolute,  given  by  Linnaeus  to  the  kind  of  vernation  in 
which  two  leaves  (conduplicate  ones  in  his  diagram)  are  put  together  so 
that  one  half  of  each  is  exterior,  the  other  interior.  That  is  just  the  mode 
in  question  reduced  to  a  single  pair  of  leaves,  as  it  is  in  the  calyx  of  a  Poppy. 
Mirbel  is  the  only  botanist  who  has  applied  the  term  to  asstivation,  and  to  a 
circle  of  more  than  two  leaves,  and  it  has  never  been  adopted  in  botanical 
descriptions.  It  has  the  disadvantage  that  the  prefix  ob  to  botanical  terms 
means  obversely  or  inversely.  Contorted  (contorta),  in  English  Tmsted,  is  in 
early  and  is  the  commonest  use,  and  it  is  sometimes  expressive.  The  objection 
to  it  is,  that  contortion  or  twisting  of  the  flower-bud  often  conspicuously  oc- 
curs where  there  is  no  overlapping  of  edges  (as  in  many  species  of  Ipomoea) ; 
that  really  no  twisting  accompanies  the  overlapping  in  a  majority  of  cases 
of  this  aestivation ;  and  that  when  there  is  a  twisting  it  is  not  rarely  in  the 
direction  contrary  to  the  overlapping;  so  that  the  contortion  needs  to  be 

FIG.  265.  Cross  section  of  the  extrorsely  plicate  or  plaited  tube  of  the  corolla  of 
a  Campanula  in  the  bud.  266.  Same  of  a  Convolvulus  (Calystegia),  the  plaits  convolute 
or  supervolute. 

FIG.  267.  Upper  part  of  unexpanded  corolla  of  Datura;  the  plaits  convolute  or 
supervolute.  268.  Cross  section  of  the  same. 


140  PHYLLOTAXY,  OR  LEAF- ARRANGEMENT. 

contorted  in  the  opposite  direction ;  that  is,  the  plaits  overlap 
to  the  right  and  are  twisted  to  the  left.1 

261.  Direction  of  Overlapping,  &c.  This  is  to  be  noted  in  the 
ternary,  quinary,  or  other  forms  of  spirally  disposed  imbrication, 
also  in  convolute  and  twisted  or  contorted  aestivation.  It 
may  be  either  to  the  right  (dextrorse)  or  to  the  left  (sinistrorse) . 
The  application  of  this  term  depends  upon  the  assumed  position 
of  the  observer,  whether  outside  or  inside.  We  always  suppose 
him  to  stand  outside,  in  front  of  the  object :  so  when  the  over- 
lapping is  from  right  to  left  of  the  observer  thus  placed,  as  in 
Fig.  266,  it  is  sinistrorse ;  when  from  his  left  to  right,  as  in 
Fig.  267,  268,  dextrorse*  The  direction  is  generally  constant, 
but  in  many  cases  only  prevalent,  in  the  same  plant  or  the  same 
species,  or  even  the  same  genus :  sometimes  it  is  uniform  or 
nearly  so  throughout  a  whole  natural  order. 


separately  expressed.  To  describe  the  aestivation  in  such  cases  as  dextrorsum 
contorta  et  sinistrarsumtorta  (or  in  similar  English  words),  when  the  overlapping 
is  to  the  right  and  the  twisting  to  the  left,  is  at  least  awkward  and  cumbrous. 
Convolute  is  a  fitting  name,  of  occasional  early  application  to  this  aestivation 
(as  by  Jussieu  to  the  petals  of  Malvaviscus),  but  without  definition  in  this 
sense ;  it  has  for  many  years  been  steadily  adopted  by  the  present  writer, 
is  employed  by  Eichler  in  Germany,  and  has  recently  been  adopted  by 
G.  Henslow  and  others  in  Great  Britain.  It  has,  however,  the  disadvantage 
of  having  been  used  by  Linnaeus  to  express  the  coiling  of  single  leaves,  and 
in  a  manner  not  wholly  congruous,  but  still  with  one  edge  outside  and  the 
other  inside. 

1  In  our  phraseology,  dextrorsely  convolute  and  sinistrorsely  contorted  ; 
in  the  current  phraseology  above  referred  to,  dextrorsely  contorted  or  twisted 
and  sinistrorsely  twisted! 

2  The  reasons  for  adopting  this  view  (in  opposition  to  the  authority  of 
Linnaeus  and  DeCandolle)  are  given  in  note  on  p.  51. 


ANTHOTAXY,   OR  INFLORESCENCE. 


141 


CHAPTER  V. 


ANTHOTAXY,   OR  INFLORESCENCE. 

262.  INFLORESCENCE,  a  term  which  would  literally  denote  the 
time  of  flower-bearing,  was  applied  by  Linnaeus  to  the  mode,  that 
is,  to  the  disposition  of  blossoms  on  the  axis  and  as  respects 
their  arrangement  with  regard  to  each  other.     ANTHOTAXY,  a 
name  formed  on  the  analogy  of  phyllotaxy,  and  denoting  flower- 
arrangement,  is  a  better  term.     The  subject  really  belongs  to 
ramification   (83,  14-16),  and  is  also  concerned  with  foliation 
and  with  phyllotaxy.     It  is  most  advantageously  treated  apart, 
immediately  preceding  the  study  of  the  blossom  itself. 

263.  In    and    near    the    blossom, 
both  axis  and  foliage  very  commonly 
undergo  modification,  either  abrupt  or 
gradual,  giving  rise   in  the  former  to 
Peduncles  and  Pedicels,  in  the  latter  to 
Bracts  and  Bractkts. 

264.  A  Bract  (in  Latin  Bractea)  is  a 
leaf    belonging    to    or    subtending    a 
flower-cluster,  or  subtending  a  flower, 
and  differing  from  the  ordinary  leaves 
in  some  respect,  usually  in  shape  and 
size,  not  rarely  in  texture  and  color.1 
They  are  commonly,  but   not  always, 
reduced  or  as  if  depauperate  leaves,  of 
little  or  no  account  as  foliage,  but  some- 
times of  use  for  protection,  sometimes 
rivalling  the  highly  colored  flower-leaves 
for  show,  more   often  insignificant  or 

minute  and  functionless,  sometimes  obsolete  (as  in  Cruciferae) ,  or 

1  Bracts  of  the  first  order  are  sometimes  called  floral  leaves  (Folia  floralia), 
or  at  least  these  are  not  well  distinguished  from  bracts.  But  the  term  floral 
leaves  is  descriptively  more  properly  and  usually  applied  to  leaves  below 
the  bracts  or  proper  origin  of  the  flower-clusters,  yet  near  them,  and  un- 
like the  proper  cauline  leaves.  It  is  a  vague  term,  and  is  in  some  danger  of 
being  confounded  (as  it  never  should  be)  with  another  vague  term,  viz. 
flower-leaves,  or  the  leaf-like  organs  of  the  flower  itself. 

FIG.  269.    Bract  (spathe)  of  Indian  Turnip,  partly  cut  away  below  to  show  the 

fleshy  spike  (spadix)  of  flowers  which  it  surrounds. 


142 


ANTHOTAXY,   OR   INFLORESCENCE. 


fugacious.  Each  flower  is  subtended  by  (grows  from  the  axil 
of)  a  bract  in  Fig.  277-280,  &c.  A  cluster  of  flowers  is  sub- 
tended by  a  conspicuous  and  colored  bract  in  Fig.  269,  270,  271 ; 
by  a  circle  of  colored  bracts,  imitating  white  petals,  in  Fig.  294. 
SPATHE  is  the  name  given  to  such  an  enclosing  bract,  or 
to  two  or  more  leaves  successively  enclosing  a  flower-cluster. 


INVOLUCRE  is  the  name  given  to  a  circle  or  spiral  collection  of 
bracts  around  a  flower-cluster,  as  in  Cornel  (Fig.  294,  also  in  Fig. 
280  and  286),  or  around  a  single  flower,  as  in  Hepatica  and 
Mallow.  A  compound  inflorescence  may  have  both  a  general 
and  a  partial  involucre,  one  for  the  general  flower-cluster,  others 
subtending  the  partial  clusters.  The  name  of  involucre  is  then 
reserved  for  the  general  one  ;  that  of 

INVOLUCEL  is  applied  to  the  partial,  secondary ,' or  ultimate 
involucres. 

BRACTLETS  (Lat.  Bracteola,  diminutive  of  bract)  are  bracts  of 
9  secondary  or  ultimate  order.  For  example,  in  the  slender 
flower-cluster,  Fig.  277,  b  is  a  bract,  subtending  each  individual 
flower-stalk  ;  b'  is  a  bractlet,  or  bract  of  secondary  order,  borne 
on  that  partial  flower-stalk  itself.  The  French  naturally  translate 
the  Latin  bracteola  into  bracteole  (pi.  bracteoles)  :  in  English, 
bractlet  is  an  idiomatic  and  better  diminutive. 

PALETS   (Lat.   Palece),  also  called   Chaff,  are  diminutive  or 

FIG.  270.  Monophyllous  spathe  of  Indian  Turnip,  with  tip  more  erect.  271.  Spatlie 
ind  spadix  of  Calla.  272.  Raceme  of  Cherry,  leafy  at  base.  273.  Dichotomous  cyme. 
Z74.  Panicle  of  Meadow-Grass.  275.  A  corymb. 


BRACTS   AND   FLOWER-STALKS.  143 

chaff-like  bracts  or  bractlets  on  the  axis  (or  receptacle)  and 
among  the  flowers  of  a  dense  inflorescence,  such  as  a  head  of 
Composite  (275,  Fig.  287,  288)  ;  and  the  name  is  also  given 
to  an  inner  series  of  the 

GLUMES  of  Grasses.  These  are  peculiar  chaffy  bracts  or  bract- 
lets  which  characterize  the  inflorescence  of  Grasses  and  Sedges. 

265.  Peduncle  is  the  general  name  of  a  flower-stalk,  that  is, 
of  an  axis  or  stem,  which  instead  of  foliage,  or  at  least  ordi- 
nary foliage,  supports  a 
flower-cluster  or  a  single 
flower.  In  Fig.  276, 
each  peduncle  (rising 
from  the  axil  of  an  ordi- 
nary leaf,  and  therefore 
answering  to  a  branch) 
bears  a  solitary  flower.  In  Fig.  277,  the  peduncle  bears  a  series 
of  flowers,  or  a  flower-cluster.  In  this  instance,  each  flower  is 
borne  on  a  flower-stalk  of  its  own,  that  is,  upon  a 

PEDICEL.  This  is  the  name  given  to  distinguish 
a  partial  flower-stalk,  or,  more  strictly,  the  stalk 
of  each  individual  flower  of  an  inflorescence.  (Fig. 
277-284.)  In  less  simple  flower-clusters,  with 
ramification  of  two,  three,  or  more  grades,  general 
peduncle,  partial  peduncles,  and  pedicels  have  to  be 
distinguished  :  the  term  pedicel  is  reserved  for  the 
ultimate  ramification. 

SCAPE  is  the  name  given  to  a  peduncle  rising 
from  the  ground,  as  that  of  most  Primulas,  of 
Dodecatheon,  Hepatica,  and  the  so-called  acaules- 
cent  or  stemless  Violets. 

RHACHIS  (backbone)  is  a  name  given  to  the  axis  » 
of  inflorescence ;  that  is,  the  continuation  of  the 
stem  or  peduncle  through  a  somewhat  elongated 
flower-cluster,  as  in  a  spike  of  Birch  or  of  Plan- 
tain, Fig.  289,  290.  When  this  axis  is  short,  as  in 
a  head  (Fig.  285-288),  it  is  usually  called  the 
RECEPTACLE,  a  word  also  used  for  the  axis  or  cauline- 
part  of  a  flower.  The  context  should  show  when  receptacle  of 
inflorescence,  and  when  receptacle  of  the  flower  itself,  is  meant. 
Both  belong  to  axis  or  stem. 

FIG.  276.  Moneywort,  Lysimachia  nummularia,  with  axillary  one-flowered 
peduncles. 

FIG.  277.  A  Raceme,  with  a  general  peduncle  (p),  pedicels  (p*),  bracts  (6),  and 
bractlets  (60- 


144  ANTHOTAXY,  OR   INFLORESCENCE. 

266.  Position   of   Flowers   or   Clusters.     Flower-buds   accord 
with  leaf-buds  in  origin,  position,  and  structure,  to  this  extent  at 
least,  that  the  parts  of  both  are  leaves  or  homologues  of  leaves, 
crowded  in  whorls  or  spirals  upon  a  short  portion  of  stem  or 
axis ;  and  as  leaf-buds  are  either  terminal  or  axillary  (15,  75) 
so  also  are  flower-buds  ;  as  a  leaf-bud  may  give  rise  to  a  simple 
or  a  compound  growth,  i.  e.  may  branch  again  and  again,  or  not 
branch  at  all,  so  flower-bearing  branches,  or  the  flower-bearing 
extremity  of  a  stem  or  branch,  may  bear  a  single  flower,  or  a 
more  or  less  compound  cluster.     Thus,  in  Fig.  276,  an  axillary 
peduncle,  or  naked  branch,  bears  at  apex  a  solitary  flower ;  in 
Fig.  277,  a  peduncle  bears  a  loose  cluster  of  flowers,  each  of 
which  springs  from  the  axil  of  a  small  bract;   in  Fig.   285, 
a  terminal  peduncle  bears   at  summit   a  dense  flower-cluster. 
Flowers  are  either  solitary  or  in  clusters.     When  solitar}-,  they 
are  naturally  without  bracts,  being  subtended  instead  (as  in  Fig. 
276)  by  ordinary  foliage. 

267.  The  elevation  either  of  a  solitary  flower  or  a  cluster  on  a 
peduncle,  or  of  individual  flowers  of  a  cluster  on  pedicels,  is  only 
incidental.    The  flowers  may  be  stalkless,  i.  e.  sessile. 

268.  The  Kinds  of  Inflorescence  which  have  received  distinctive 
names  are  various,  but  are  all  reducible  to  two  types,  which, 
generally  well  marked,  may  sometimes  pass  into  each  other,  and 
which  are  not  rarely  combined  in  the  same  compound  inflores- 
cence.1   The  two  types  differ  in  basis  as  do  axillary  from  ter- 
minal buds  ;  in  the  one  the  flowers  are  axillary  or  lateral,  in  the 
other  terminal  in  respect  to  the  axis  from  which  each  flower  01 
its  pedicel  arises.     But  inasmuch  as  every  flower,  whatever  its 
position,  is  terminal  to  its  own  stalk  or  axis,  it  is  better  to  dis- 
tinguish the  two  types  in  other  terms,  and  to  name  them  the 

269.  Indefinite  and  Definite,  or,  in  equivalent  and  similar  terms, 
the  Indeterminate  and  Determinate.2     Each  may  be  either  simple 
or  compound.     It  is  from  the  simple  that  the  definitions  are  to 
be  drawn.     In  the  former  type,  the  rhachis  or  main  axis  of  the 
inflorescence  is  not  terminated  by  a  flower,  but  lateral  axes,  or 
pedicels,  are.     In  the  latter,  both  the  main  or  primary  and  the 
lateral  or  secondary  axes  or  stalks  are  so  terminated.     An  inde- 
terminate flower-cluster  may  go  on  to  develop  internode  after 

1  Inflorescence,  as  has  been  well  insisted  on  by  Guillaud  (in  Bull.  Soc. 
Bot.  France,  iv.  29),  is  a  mode,  not  a  thing.  The  things  sometimes  but  in- 
appropriately so  called  are  flower-clusters,  for  which,  if  a  general  technical 
name  is  needed,  that  otAnthemia,  in  English  Anthemy,  suggested  by  Guillaud, 
is  as  good  as  any. 

a  Also  named  by  Eichler  (Bliithendiagramme,  33,  following  Guillaud,  1.  c.) 
the  Cymo»e  and  the  Botryose  type. 


THE   TWO   TYPES. 


145 


internode  of  axis,  and  one  or  more  leaves  (bracts)  at  each  node, 
and  then  a  flower  in  the  axil  of  each  bract,  until  its  strength  or 
capability  is  exhausted.  Or  it  may  stop  short  with  very  few 
flowers  ;  but  the  uppermost  and  youngest  one  will  not  really  ter- 


minate  the  rhachis  (».  e.  come  from  a  terminal  bud) ,  though  it 
may  appear  to  do  so.  (Fig.  272,  277-279,  &c.)  The  lower 
flower-buds  are  evidently  the  oldest,  and  accordingly  the  first  to 
expand ;  and  the  expansion  will  proceed  regularly  from  below 
upward :  wherefore  this  type  of  inflorescence  has  been  called  the 
Ascending  or  Acropetal ;  likewise  the  Centripetal,  because,  when 
the  flowers  are  brought  to  the  same  level  or  near  it  (as  in  Fig. 
279,  280)  by  a  lengthening  of  the  lower  pedicels,  with  or  with- 
out relative  shortening  of  the  rhachis,  the  evolution 
is  seen  to  proceed  from  circumference  to  centre. 
There  is  thus  no  lack  of  names ;  but,  inasmuch  as 
\  r  the  following  type  is  commonly  referred  to  under  the 
^*  general  name  of  Cymose,  to  this  has  recently  been 
given  the  counterpart  name  of  Botryose.  (271.) 


«l 


VfljL 


\ 


281  282  283  284 

270.  A  determinate  flower-cluster  (as  seen  in  its  gradual 
development  which  is  not  rarely  presented)  has  the  last  internode 

FIG.  278-280.  Diagrams  of  indefinite,  indeterminate,  contripetal,  or  botryose  in- 
florescence :  278,  Raceme ;  279,  Corymb ;  280,  Umbel. 

FIG.  281-284.  Diagrams  of  definite,  determinate,  centrifugal,  or  cymose  inflores- 
cence :  281,  a  false  or  descending  raceme ;  282,  a  solitary  terminal  flower ;  283,  same  with 
two  lateral  flowers  developing,  forming  a  3-flowered  cyme;  284,  same  with  lateral 
peduncles  3-flowered,  or  a  pair  of  3-flowered  cymes,  beside  the  central  or  primary 
terminal  flower. 


146  ANTHOTAXY,  OR   INFLORESCENCE. 

of  its  axis  terminated  by  a  flower  (Fig.  281-284) ,  which  answers 
to  a  terminal  bud.  If  more  flowers  appear,  so  as  to  compose  a 
cluster,  they  spring  from  the  axils,  preferably  from  the  highest 
axils,  and  are  later.  The  order  of  evolution  is  shown  in  the  figures 
by  the  size  of  the  flower-buds  or  degree  of  expansion  of  the 
blossoms.  Fig.  281  best  shows  why  a  determinate  or  definite  in- 
florescence is  sometimes  said  to  be  Descending ;  Fig.  283  shows 
why  it  is  called  Centrifugal,  the  central  flower  first  expanding  ; 
Fig.  284  exhibits  the  lateral  or  circumferential  partial  clusters 
later  than  the  central  blossom,  and  their  lateral  flowers  later 
than  their  central. 

271.  Varieties  of  Indeterminate  or  Botryose  Inflorescence.     The 
names  of  most  of  these  have  been  fixed  from  the  time  of  Linnaeus, 
but  defined  without  reference  to  the  order  of  evolution  of  the 
flowers.     They  are  the  Raceme,  Corymb,  and  Umbel,  with  flowers 
raised  on  pedicels ;  the  Spike  and  Head,  with  sessile  flowers ; 
also  some  modifications  of  these,  notably  the  Ament  and  the 
Spadix.     The  raceme  may  be   taken  as   the  type.     Botrys  is 
equivalent  to  racemus,  &c.  ;  and,  as  the  type  includes  diversity 
of  forms  to  which  the  name  racemose  would  seem  inapplicable, 
the  term  botryose  (botrytischen  of  Eichler)  is  best  chosen  as  the 
general  name  of  it,  and  is  a  good  counterpart  to  cymose  for  the 
other  type. 

272.  A  Raceme  (illustrated  in  Fig.  272,  277,  and  by  diagram 
in  Fig.  278)  is  a  simple  flower-cluster,  in  which  the  flowers, 
on  their  own  lateral  or  axillary  pedicels  and  of  somewhat  equal 
length,  are  ai-ranged  along  a  relatively  more  or  less  elongated 
rhachis  or  axis  of  inflorescence.     The  common  Barberry,  Cur- 
rant, Choke-Cherry  and  Black  Cherry,  and  Lily  of  the  Valley 
are  familiar  examples. 

273.  A  Corymb    (Fig.  275,    279)   is   a   shorter  and  broader 
botr3rose  cluster,  which  differs  from  a  raceme  only  in  the  relatively 
shorter  rhachis  and  longer  lower  pedicels ;  the  cluster  thus  be- 
coming flat-topped  or  convex.     The  centripetal  character  is  thus 
made  apparent.     The  greater  number  of  the  corymbs  of  Linnasus 
and  succeeding  botanists  are  cymes,  the  central  flower  first  ex- 
panding.    And  the  term  corymbose  or  corymb-like  is  still  much 
used  in  descriptive  botany  for  a  ramification  which  is  mainly  of 
the  cymose  type,  and  where  in  strictness  the  term  cymose  should 
be  employed. 

274.  An  Umbel  (Fig.  280),  as  in  Asclepias,  &c.,  differs  from 
a  corymb  only  in  the  extreme  abbreviation  of  the  rhachis  or  axis 
of  inflorescence,  and  the  general  equality  of  the  pedicels  which 
thus  all  appear  to  originate  from  the  apex  of  the  peduncle,  and 


THE  BOTKYOSE   TYPE. 


147 


so  resemble  the  rays  of  an  umbrella ;  whence  the  name,  and 
whence  also  the  pedicels  or  partial  peduncles  of  an  umbel  are 
termed  its  Rays.  The  bracts,  brought  by  the  non-development 
of  internodes  into  a  depressed  spiral  or  apparent  (or  sometimes 
real)  whorl,  become  an  involucre.  (264.)  An  umbel  or  any 
similar  cluster  when  sessile  (without  a  common  peduncle) ,  and 
the  parts  crowded,  is  sometimes  called  a  Fascicle  (or  the  pedicels 
said  to  be  fascicled)  ;  but  this  term  has  been  differently  denned. 
(280.)  It  is  better  not  to  use  it  for  any  special  kind  of  inflores- 
cence, but  simply  in  the  sense  of  a  bundle  of  whatever  sort. 
This  will  accord  with  the  sense  in  which  it  is  applied  to  an 
aggregation  of  leaves.  (248.) 

275.  A  Head  or  Capitulum  (Fig.  285)  is  a  globular  cluster  of 
sessile  flowers,  like  those  of  Red  Clover,  Button-bush,  and 
Plane-tree.  The  pedicels  need  not  be  absolutely  wanting,  but 
only  very  short.  An  umbel 
with  pedicels  much  abbrevi- 
ated thus  passes  into  a  head, 
as  in  Eryngium,  &c.  And 
a  head  with  rhachis  elongated 
passes  into  a  spike.  The 
short  rhachis  of  a  head  very 
commonly  takes  the  name  of 
receptacle.  (265.)  The  whole 
may  be  subtended  by  con- 
spicuous bracts  forming  an 
involucre  (264)  as  in  Fig. 
286,  or  may  be  destitute  of 
any,  as  in  Fig.  285.  On  ac- 
count of  the  compactness 
and  mutual  pressure  under 
growth,  the  bracts  among 
the  flowers  of  such  heads  (normally  one  subtending  each  blos- 
som) are  apt  to  be  rudimentary,  reduced  to  little  scales,  or 
abortive,  or  completely  wanting.  In  the  latter  case,  the  recep- 
tacle is  said  to  be  naked  (nude) ,  i.  e.  naked  of  bracts :  when 
they  are  present,  it  is  paleate  or  chaffy.  A  peculiar  sort  of  head, 
not  undeserving  a  special  name  (though  this  is  not  necessary 
in  descriptive  botany) ,  is  the 

ANTHODIUM,  the  so-called  Compound  Flower  of  the  earlier 
botanists,  which  gives  the  name  to  the  vast  order  of  Composit*. 


FIG.  285.    Cephalanthus  occideiitalis,  the  Button-bush;  a  pair  of  leaves,  and  a 
terminal  peduncle  bearing  a  dense  head  (capitulum)  of  flowers. 


148  ANTHOTAXY,  OR   INFLORESCENCE. 

The  name  means  "resembling  a  flower."    Although  it  has  all 
the  characters  of  a  true  head,  the  resemblance  to  a  flower  is 


remarkably  striking,  the  involucre  imitating  a  calyx,  and  the 
strap-shaped  (ligulate)  corollas  of  the  several  flowers  imitating 
the  petals  of  a  single  blossom.  In  some  (such  as  Dandelion 

and  the  Cichory,  Fig.  286) ,  all 
the  flowers  of  the  head  bear 
these  petal-like  corollas;  in 
more  (such  as  Aster,  Sun- 
flower, and  Coreopsis,  Fig. 
287),  only  an  outer  circle  of 
flowers  does  so  ;  the  remain- 
der, smaller  and  filling  the 
centre  (or  disk),  may  by  the 
casual  observer  be  taken  for 
stamens  and  pistils,  and  further  the  deception.  The  rhachis 
or  receptacle  of  a  head  of  this  kind  is  commonly  depressed, 
bearing  the  flowers  on  what  then  becomes  the  upper  surface, 
which  adds  to  the  imitation.1 

SYCONIUM.  This  name,  given  to  the  Fig-fruit,  should  be  here 
referred  to,  as  it  is  a  sort  of  inflorescence,  of  the  general  nature 
of  a  head,  but  with  receptacle  external  and  flowers  enclosed 

1  The  receptacle  of  an  Anthodium  has  been  termed  Clinanthium  or  Phor- 
anthium ;  and  its  involucre,  a  Periphoranthium  or  Periclinium.  The  head  has 
likewise  been  named  a  Cephalanthium. 

FIG.  286.  Flowering  branch  of  Cichory,  with  two  heads  of  ligulate  flowers,  of 
natural  size. 

FIG.  287.    Vertical  section  of  a  head  of  flowers  of  a  Coreopsis. 


THE   BOTEYOSE   TYPE. 


149 


within.  See  Fig.  657-659  (683) ,  where  its  morphology  is  ex- 
plained and  illustrated.  Viewed  as  an  inflorescence,  it  has  also 
been  named  a  HYPANTHODIUM. 


276.  A  Spike  is  a  cluster  of  sessile  (or  apparently  or  nearly 
sessile)  lateral  flowers  on  an  elongated  axis.  It  may  be  de- 
fined by  comparison,  as  a  head  with  the  rhachis  lengthened 
(indeed  a  young  head  often  becomes  a  spike  when  older) ,  and 
equally  as  a  raceme  with  the  pedicels 
all  much  shortened  or  wanting.  A 
common  Mullein  and  a  Plantain 
(Plantago,  Fig.  290)  are  familiar  ex- 
amples. Two  modifications  of  the 
spike  (or  sometimes  of  the  head)  gen- 
erally bear  distinct  names,  although 
not  distinguishable  by  exact  and  con- 
stant characters,  viz. :  — 

SPADIX,  a  spike  or  head  with  a 
fleshy  or  thickened  rhachis.  The 
term  is  almost  restricted  to  the  Arum 
family  and  Palms,  and  to  cases  in 
which  the  inflorescence  is  accompanied 
by  the  peculiar  bract  or  bracts  called 
a  spathe  (Fig.  269-271).  But  the 
two  do  not  always  go  together:  in 
Acorus  and  Orontium  there  is  properly 
no  spathe  to  the  spadix  ;  while  in  the 
Iris  family  the  bracts  are  said  to  form 
a  spathe,  and  there  is  no  spadix.  In  Palms,  the  principal  reason 
for  naming  the  inflorescence  a  spadix  is  its  inclusion  in  a 
spathe  before  anthesis. 

PIG.  288.  A  slice  of  Fig.  287,  more  enlarged,  with  one  tubular  perfect  flower  (a)  left 
standing  on  the  receptacle,  and  subtended  by  its  bract  or  chaff  (6);  also  oneligulate 
and  neutral  ray-flower  and  part  of  another  (c,  c) :  in  d,  d,  the  bracts  or  leaves  of  the 
involucre  are  seen  in  section. 

PIG.  289.    Catkin  of  White  Birch.    290.  Young  spike  of  Plantago  major. 


150 


ANTHOTAXY,  OR  INFLORESCENCE. 


AMENT  or  CATKIN.  This  is  merely  that  kind  of  spike  with 
scaly  bracts  borne  by  the  Birch  (Fig.  289),  Poplar,  Willow,  and, 
as  to  one  sort  of  flowers,  by  the  Oak,  Walnut,  and  Hickory, 
which  are  accordingly  called  amentaceous  trees.  Catkins  usually 
fall  off  in  one  piece,  after  flowering  or  fruiting.  All  true  catkins 
are  unisexual. 

277.  Any  of  these  forms  of  simple  inflorescence  may  be  com- 
pounded. Racemes  may  themselves  be  disposed  in  racemes,  spikes 
in  spikes  (as  in  Triticum) ,  heads 
be  aggregated  in  heads,  umbels  in 
umbels,  corymbs  may  be  corym- 
bosely  compound,  &c. ;  forming 
compound  racemes,  spikes,  umbels, 
and  the  like,  the  terminology  of 
which  is  easy.  The  most  usual 
case  of  truly  homomorphous  com- 
pounding is  that  of  umbels  ;  the 

Inflorescence  of  much  the  larger  part  of  Umbelliferse  being  in 
compound  umbels,  as  in  Fig.  290°.  There  is  then  the  general 
umbel,  the  rays  of  which  become  peduncles  to 
the  partial  umbels,  and  the  rays  of  the  latter 
are  pedicels.  Umbella  and  Umbellulce  desig- 
nate in  Latin  terminology  the  general  and  its 
partial  umbels.  Umbellets  (coined  by  the  late 
Dr.  Darlington)  may  well  replace  the  latter 
as  the  English  diminutive.  But  umbels  are 
sometimes  racemose!}'  arranged,  as  in  Aralia 
spinosa,  heads  may  be  arranged  in  spikes, 
and  so  on. 

278.  A  Panicle,  of  the  simple  and  normal 
sort  (as  illustrated  in  Fig.  291),  is  produced 
when  a  raceme  becomes  irregularly  compound 
by  some  (usually  the  lower)  of  its  pedicels 
developing  into  peduncles  carrying  several 
flowers,  or  more  than  one,  or  branching  again 
and  again  in  the  same  order.  But  in  com- 
pound clusters  generally  the  secondary  and 
tertiary  ramifications  are  apt  to  differ  in  t}-pe 
as  well  as  in  particular  mode,  giving  rise  to 
heteromorphous  or  mixed  inflorescence.  (288.) 
As  Linnaeus  defined  the  term,  and  as  it  has  generally  been  em- 
ployed in  botanical  descriptions,  the  panicle  is  a  general  term 


FIG.  290».    Compound  umbel  of  Caraway.    291.    A  simple  panicle. 


THE   CYMOSE   TYPE.  151 

for  any  loose  and  diversely  branched  cluster,  with  pedicellate 
flowers.  It  is  therefore  difficult  to  restrict  it  in  practice  to  the 
indeterminate  type. 

279.  Varieties  of  Determinate  or  Cymose  Inflorescence.     The 
plan  of  this  type  has  been  sufficiently  explained.    (270.)     Its 
simplest  condition  is  that  of  a  solitary  terminal  flower,  peduncu- 
late or  pedicellate  (as  in  Fig.  282),  or  sessile.     The  production 
of  more  flowers  necessitates  new  axes  from  beneath,  from  the 
axils  of  adjacent  leaves  or  bracts.     These,  being  later,  render 
the  evolution  centrifugal.     The  simplest  flower-cluster  (unless 
we  call  the  solitary  flower  of  Fig.  282  a  one-flowered  cluster)  is 
that  of  Fig.  283,  where  a  secondary  floral  axis  or  peduncle  has 
developed  from  the  axil  of  each  leaf  of  the  uppermost  pair,  or 
where  with  alternate  leaves  there  is  a  single  uppermost  leaf,  and 
then  only  one  such  peduncle,  and  thus  is  produced  a  three-  (or 
two-)  flowered  cymose  cluster.     The  flower  of  the  primary  axis 
is  marked  by  its  bractless  peduncle  (therefore  a  pedicel)  ;  the 
lateral  and  secondary  peduncles  are  known  (commonly  or  nor- 
mally) by  their  bracts  or  bract ;  the  portion  below  the  bracts 
is  proper  peduncle ;  that  above,  of  single   internode,   pedicel. 
Bracts,  like  other  leaves,  have  potential  buds  in  their  axils  ;  these 
in   an  inflorescence  give   the  third  order  of  ramification,  each 
branch  tipped  with  its  flower ;  and  so  on. 

280.  The  Cyme  is  the  general  name  of  this  kind  of  flower-cluster 
in  its  various  forms.     One  of  these  very  simple  cjnnes,  by  itself  or 
as  a  part  of  a  larger  cyme,  may  be  called  a  Cymule.     The  regular 


cyme  usually  accompanies  opposite  or  other  grades  of  verticillate 
leaves,  but  is  not  rare  in  the  alternate  arrangement.  It  is 
readiest  understood  in  an  opposite-leaved  plant  with  regular 
opposite  ramification,  as  in  an  Arenaria,  Fig.  292.  By  its  con- 
stitution, a  cyme  proceeds  from  simple  to  compound.  It  mat- 

PIG.  292.    Dichotomous  or  biparous  cyme  (cyme  bipare  of  Bravais,  Dichasium  of 
Eichler)  of  Arenaria  Michauxii. 


152  ANTHOTAXY,  OB   INFLORESCENCE. 

ters  little  whether  its  development  is  progressive,  the  flowers  of 
the  ultimate  ramifications  expanding  after  the  earlier  have  matured 
fruit,  and  with  subtending  bracts  conspicuous  or  foliaceous ;  or 
whether,  as  in  Elder  and  Hydrangea  (Fig.  293,  and  in  Fig. 
273),  the  bracts  are  minute  and  caducous  or  abortive,  and  the 
ramification  complete  with  all  the  flower-buds  well  formed  before 
the  oldest  expand,  so  that  the  whole  is  in  blossom  almost  at  the 
same  time.  But  a  cyme  may  be  properly  said  to  be  compound 
when  the  primary  axis  in  it  is  a  peduncle  instead  of  a  pedicel, 


and  supports  a  cluster  (cyme  or  cymule)  instead  of  a  solitary 
central  flower  at  the  main  divisions.1  One  form  of  the  regular 
cyme,  on  account  of  its  compactness,  is  named  the 

GLOMERULE.  This  is  merely  a  cymose  inflorescence,  of  any 
sort,  which  is  condensed  into  the  form  of  a  head,  or  approach- 
ing it.  Of  this  kind  is  the  so-called  head  of  Cornus  florida,  and 
of  the  herbaceous  C.  Canadensis  (Fig.  294),  which  shows  the 

1  The  dichotomous  or  two-branched  cyme  is  the  commonest,  but  is  some- 
times marked  by  suppression  of  internodes ;  as,  for  example,  where  the 
branches  are  apparently  in  fours,  in  an  umbelliform  way ;  but  these  are  two 
sets  of  two,  with  the  internode  between  the  pairs  extremely  short ;  or  where, 
as  in  Elder,  the  branches  or  rays  are  five,  in  this  case  consisting  of  the  same 
two  pairs  and  a  central  one,  which  is  a  many-flowered  continuation  of  the 
primary  axis.  Or  5-rayed  cymes,  &c.,  may  be  founded  upon  alternate  leaves 
with  shortened  internodes,  the  rays  or  peduncles  axillary  to  them  thus 
brought  into  an  apparent  whorl. 

Bravais  distinguished  cymes  as  mvltiparous,  with  three  or  more  lateral 
axes ;  biparous,  with  two ;  and  uniparous,  with  only  one  (cyme  multipare, 
bipare,  unipare).  To  these  Eichler  gives  the  substantive  names,  severally, 
of  Pleiochasium,  Dichasium,  and  Monochasium.  Only  the  latter  needs  illustra- 
tion ;  the  others  being  as  it  were  compounds  of  this. 

FIG.  293.  Compound  cyme  of  Hydrangea ;  with  some  neutral  and  enlarged  mar- 
ginal flowers. 


THE  CYMOSE  TYPE. 


153 


composition  best,  on  close  examination.  A  condensed  but  less 
capitate  cyme,  or  cluster  of  cymes,  was  called  by  Roeper  and 
DeCandolle  a  FASCICLE  ;  and  this  terminology  has  been  much 
adopted.  It  is  properly  enough 
said  to  be  a  fascicle,  which,  as 
used  by  Linnaeus  and  others, 
means  a  bundle,  or  close  collection 
of  parts,  whether  leaves,  pedun- 
cles, or  flowers ;  but  a  fascicle  is 
not  necessarily  a  cyme  (274) ,  nor  is 
there  need  of  a  special  substantive 
name  for  a  compact  cyme,  which 
may  either  be  simply  so  called  or 
it  may  pass  into  the  glomerule. 

281.  Botryoidal  forms  of  Cymose 
Type,  or  False  Racemes,  &c.  The 
regular  cyme  seldom  continues 
with  all  its  ramifications.  In 
Fig.  292,  after  the  second  forking, 
one  of  the  two  lateral  peduncles 
mostly  fails  to  appear,  and  in  some 
parts  one  of  the  bracts  also  ;  and  ultimately  the  lateral  peduncle 
present  is  bractless,  like  the  central,  therefore  equally  incapable 
of  further  ramification,  being  reduced  to  a  pedicel  of  a  single  inter- 
node.  This  suppression  some- 
times begins  at  the  first  fork- 
ing or  at  the  very  base  ;  and, 
when  followed  throughout,  it 
reduces  a  biparous  or  dichoto- 
mous  cyme  to  one  half,  and, 
converts  this  half  (when  the 
axis  straightens)  into  the  sem- 
blance of  a  raceme  if  the 
flowers  are  pedicelled,  or  of  a 
spike  when  they  are  sessile. 
Fig.  296  is  a  diagram  of  such  an  inflorescence  as  that  of  Fig.  292, 
with  one  lateral  branch  uniformly  suppressed  at  each  division,  the 
wanting  members  indicated  by  short  dotted  lines.  Cases  exem- 
plifying this  occur  in  portions  of  the  inflorescence  of  some  of  our 

FIG.  294.  Plant  of  Cornus  Canadensis :  flowering  stem  bearing  a  cluster  of  leaves 
above,  then  continued  into  a  peduncle,  and  terminated  by  a  glomerule  of  very  small 
flowers;  this  subtended  by  a  colored  and  corolla-like  involucre  of  four  bracts.  295.  One 
of  the  flowers  taken  from  the  glomerule,  enlarged. 

FIG.  296.    Uniparous  cyme  or  sympodial  false  raceme,  with  opposite  leaves  or  bracts. 

FIG.  297.    Form  of  the  same,  with  alternate  leaves  or  bracts. 


154  ANTHOTAXY,  OR   INFLORESCENCE. 

smaller  Hypericums,  and  notably  in  H.  Sarothra,  in  which  the 
leaves  are  all  reduced  to  bracts.  It  is  not  alwaj-s  easy  to  show 
why  this  is  not  a  true  raceme .  But  the  other  bract  of  the  pair,  upon 
that  supposition,  is  unaccountably  empty  :  the  successive  angular 
divergence  of  each  joint  of  the  axis  of  inflorescence  in  the  younger 
part,  which  commonly  runs  into  a  coil,  finds  explanation  in  the  view 
that  each  portion  is  the  lateral  branch  from  the  axil  of  the  subtend- 
ing leaf:  and  occasionally  the  other  axil  produces  a  similar  one, 
thus  revealing  the  cymose  character.  When  the  bract  from  the 
axil  of  which  the  missing  branch  should  come  disappears  also,  as 
sometimes  it  does,  and  uniformly  on  the  same  side,  a  state  of 
things  like  that  of  the  upper  part  of  Fig.  297  occurs.  The  same 
figure  may  serve  for  the  arrangement  corresponding  to  that  of 
Fig.  296,  only  with  alternate  leaves.  But  then,  close  as  the  imi- 
tation of  a  raceme  here  is,  the  position  of  each  flower  in  respect 
to  the  bract  supplies  a  criterion.  While  in  a  true  raceme  the 
flower  stands  in  the  axil  of  its  bract,  here  it  stands  on  the  oppo- 
site side  of  the  axis,  or  at  least  is  quite  away  from  the  axil. 

282.  Sympodial  forms.     The  explanation  is  that  the  axis  of 
inflorescence  in  such  cases,  continuous  as  it  appears  to  be,  is 
not  a  simple  one,  is  not  a  monopode,  but  a  sympode  (110,  116, 
notes) ,  i.  e.  consists  of  a  series  of  seemingly  superposed  inter- 
nodes  which  belong  to  successive  generations  of  axes  :  each  axis 
bears  a  pair  of  leaves  (Fig.  296)  or  a  single  leaf  (Fig.  297), 
is  continued    beyond    into  a   peduncle    (or    pedicel  in  these 
instances),   and   is   terminated   by  a   flower.      From   the   axil 
promptly  springs  a  new  axis  or  branch,  vigorous  enough  soon 
to  throw   the   adjacent  pedicel  and   flower  to   one   side :  this 
bears  its  leaf  or  pair  of  leaves,  and  is  terminated  like  its  prede- 
cessor with  a  flower ;  and  so  on  indefinitely.     The  fact  that  the 
alternate  leaves  or  bracts  are  thrown  more  or  less  strictly  to  one 
side  and  the  flowers  to  the  other,  in  Fig.  297,  shows  that  these 
leaves  do  not  belong  to  one  and  the  same  axis ;  for  alternate 
leaves  are  never  one-ranked  or  disposed  preponderatingly  along 
one  side  of  an  axis,  as  in  this  diagram,  and  as  is  seen  in  the 
inflorescence  of  a  Houseleek,  &c. 

283.  A  further  difficulty  in  the  morphology  of  clusters  of  this 
class  comes  from  the  early  abortion  or  complete  suppression  of 
bracts.     This  is  not  unknown  in  botryose  inflorescence,  occurring 
in  the  racemes  of  almost  all  Cruciferae  :  it  is  very  common  in  the 
cymose  of  all  varieties,  and  especially  in  the  uniparous  ones  in 
question,  which  characterize  or  abound  in  Borraginacese,  Hydro- 
phyllaceae,  and  other  natural  orders.     In  some  genera  or  species, 
the  bracts  are  present,  or  at  least  the  lower  ones ;  in  others, 


THE   CYMOSE  TYPE.  155 

absent ;  in  some,  either  occasionally  present  or  wanting  in  the 
same  species  or  individual.  It  is  only  by  analogy,  therefore, 
and  by  a  comparison  of  allied  plants,  that  the  nature  of  some  of 
these  flower-clusters  can  be  made  out.  With  the  botanists  of  a 
preceding  generation,  these  one-sided  clusters  were  all  described 
as  racemes  or  spikes.  Botanists  still  find  it  convenient  to  con- 
tinue the  use  of  these  names  for  them  in  botanical  descriptions, 
adding,  however,  as  occasion  requires,  the  qualification  that  they 
are  false  racemes  or  spikes,  or  cymose  racemes,  and  the  like  ;  or 
else,  by  reversing  the  phrase,  with  stricter  correctness  they  call 
them  racemiform  or  spiciform  cymes,  &c. 

284.  Commonly  these  false  racemes  or  spikes  (or  botryoidal 
cymes,  if  we  so  name  them)  are  circinate  or  inrolled  from  the 
apex  when  young,  in  the  manner  of  a  crosier,  straightening  as 
they  come  into  blossom  or  fruiting.     Likening  them  to  a  scorpion 
when  coiled,  the  earlier  botanists  designated  this  as  scorpioid. 
As  the  coil  is  a  helix,  it  has  also  been  named  helicoid.1    The 
flowers  are  then  thrown,  more  or  less  strictly,  to  the  outer  side 
of  the  coiled  rhachis,  where  there  is  room  for  them  ;  and  so  these 
False  racemes  or  spikes  are  secund  or  unilateral.     The  particular 
anthotaxy  and  phyllotaxy  of  the  various  sympodial  and  botryoi- 
dal forms  of  cymose  inflorescence  become  rather  difficult ;  and 
the  sorts  which  have  been   elaborately  classified  into  species 
(and  have  no  little  morphological  interest)   are  connected  by 
such  transitions,  and  are  based  on  such  nice  or  sometimes  theo- 
retical particulars,  that  the  terminology  based  on  them  is  seldom 
conveniently  applicable  io  descriptive  botany,  at  least  as  to  sub- 
stantive names. 

285.  One  of  the  latest  and  simplest  classifications  of  cymes  is 
that  of  Eichler  in  his  Bliithendiagramme.2 

1  Scorpioid  and  Helicoid  have  been  carefully  distinguished  by  later 
morphologists,  on  account  of  some  difference  in  the  mode  of  evolution  and 
arrangement  of  the  flowers  along  one  side  of  the  rhachis,  by  which  they 
become  two-ranked  in  scorpioid,  one-ranked  in  helicoid.  But  practically 
the  two  kinds  of  clusters  are  not  always  readily  discriminated ;  and  in  gen- 
eral terminology  a  single  name,  with  subordinate  qualifying  terms,  is  suffi- 
cient. Scorpioid  is  the  older  and  commoner  one,  therefore  the  most  proper 
to  be  used  in  the  generic  sense. 
2  CYMOSE  TYPE  (classified  without  reference  to  bracts,  which  are  so  often 

wanting) ;  divided  into 
o.  Lateral  axes   three   or  more :    PLEIOCHASIUM,  the   multiparous  cyme  of 

Bravais. 

)8.  Lateral  axes  two :  DICHASIUM,  the  biparous  cyme  of  Bravais. 
y.  Lateral  axis  one  •  MONOCHASIUM,  the  uniparous  cyme  of  Bravais. 

The  latter,  or  the  corresponding  divisions  of  the  preceding  sorts,  may  be 
divided  as  follows : 


156 


ANTHOTAXY,  OR  INFLORESCENCE. 


286.  Sundry  complications  and  obscurities  are  occasionally 
encountered  in  anthotaxy  or  phyllotaxy,  which  cannot  here  be 

*  Lateral  axes  transverse  to  the  relatively  main  axis. 

1.  Lateral  axes  in   successive    generations    always   falling    on   the   same 

side  of  the  relatively  main  axis :  SCHRAUBEL  [screwlike]  or  BOSTRYX 
[ringlet  or  curl],  the  uniparous  helicoid  cyme  of  Bravais. 

2.  Lateral  axes  falling  alternately  on  opposite  sides  of  the  relatively  main 

axis:  WICKEL  or  CINCINNUS  [a  curl], the  uniparous  scorpioid  cyme  of 
Bravais. 
*  *  Lateral  axes  medial  [in  the  same  plane]  relative  to  the  main  axis. 

3.  Lateral  axes  in  successive  generations  always  on  the  back  side  of  the 

axis  from  which  it  springs:  FACHEL,  RHIPIDIUM  [fan]. 

4.  Lateral  axes  in  successive  generations  always  on  the  upper  side  of  the 

axis  from  which  it  springs  :  SICHEL,  DREPANIUM  [sickle]. 
The  subjoined  simple   diagrams  from  Eichler  (Fig.  298-303)  illustrate 
these  forms.     The  ramification  is  given  without  the  bracts,  which  theoreti- 
cally or  actually  subtend  the  axes  of  each  generation.     The  student  may  add 
them,  and  so  more  readily  apprehend  the  .characters. 

Eichler  recognizes  the  forms  with  median  (antero-posterior)  position  of 
j  axes  in  Monocotyledons  only. 

It  is  natural  to  distichous  phyl- 
lotaxy, and  it  accords  with 
the  general  rule  that,  in  mono- 
cotyledonous  plants,  the  first 
leaf  of  the  branch,  or  the  com- 
monly solitary  bractlet  of  the 
)6  peduncle,  stands  over  against 
and  facing  the  bract  or  leaf 
from  the  axil  of  which  said 
branch  or  peduncle  springs,  i.  e.  is  posterior  and  next  the  parent  axis,  as 
shown  in  the  diagram,  Fig.  304,  305. 


FIG.  298.  Diagram  of  the  Cincinnus.  299.  Diagram  of  the  Bostryx.  The  flower- 
axes  numbered  in  succession. 

FIG.  300.  Diagram  of  the  Rhipidium.  302.  Ground  plan,  indicating  the  order  of 
evolution  of  the  flowers. 

FIG.  301.  Diagram  of  the  Drepanium.  303.  Ground  plan,  the  flowers  evolved  in 
succession,  from  left  to  right. 

PIG.  304.  Diagram  showing  the  position  of  bractlet  or  first  leaf  on  a  branch  in 
Monocotyledons:  a  is  the  primary,  a/  the  secondary  axis;  b  is  bract,  and  &/  bractlet. 


THE   CYMOSE   TYPE.  157 

explained,  except  through  full  details  :  such  as  flowers  standing 
by  the  side  of  a  leaf,  or  a  small  leaf  by  the  side  of  a  larger  one, 

The  transverse  or  oblique  position  of  secondary  axes  or  peduncles,  as  in 
Eichler's  first  two  species,  brings  the  flowers  of  the  false  raceme  or  spike 
out  of  line  of  the  sympodial  axis  and  bracts,  neither  in  the  axils,  as  in  true 
racemes,  nor  opposite  them,  as  in  the  Rhipidium  and  Drepanium,  but  on  one 
side  of  this  plane  or  the  other.  This  is  most  common  in  Dicotyledons  (in 
Drosera,  Sedum,  Sempervivum,  and  Hyoscyamus,  in  Borraginaceae  and 
Hydrophyllaces,  &c.),  and  is  not  rare  in  Monocotyledons,  especially  with 
tristichous  phyllotaxy,  as  in  Tradescantia.  In  the  Bostryx,  Fig.  299,  the 
bractlet  is  anterior  or  falls  on  the  same  side  as  the  bract,  or,  in  other  terms, 
the  successive  bracts  are  all  on  one  side,  the  inner  side,  of  the  helix ;  and  the 
Drepanium  ( Fig.  301 )  is  like  it :  this  is  the  helicoid  cyme  of  Bravais,  &c., 
and  its  flowers  are  commonly  one-ranked.  In  the  Cincinnus  or  true  scorpioid 
cyme  (Fig.  298),  and  equally  in  the  Rhipidium  (Fig.  300),  the  bracts  fall 
alternately  on  opposite  or  different  sides  of  the  sympodial  rhachis,  because 
the  single  bract  (&')  of  each  successive  secondary  axis  (a')  stands  next  the 
axis  (a)  and  over  against  the  bract  (b)  of  the  generation  preceding.  The 
flowers  in  these  generally  fall  into  two  parallel  ranks  (conspicuously  so 
when  crowded)  on  the  upper  side  of  the  rhachis,  on  which,  in  the  cincinnus 
or  true  scorpioid  cluster,  they  are  usually  sessile  or  nearly  so  (or  spicate), 
as  is  well  seen  in  Heliotropes,  and  in  very  many  Borraginaceous  and  Hydro- 
phyllaceous  species,  in  Houseleek,  Tradescantia  erecta,  &c.  This  comes 
through  antidromy,  that  is,  the  phyllotaxy  of 
each  successive  axis  of  the  sympode  (with  its 
one  bract,  or  by  suppression  without  it)  changes 
direction,  from  right  to  left  and  from  left  to 
right  alternately.  Fig.  301  is  a  plan  of  this 
two-ranked  unilateral  arrangement.  When  not 
too  crowded,  both  Cincinnus  and  Rhipidium 
are  apt  to  have  a  zigzag  rhachis. 

These  two  last-mentioned  kinds  are  so  gen- 
erally alike  in  character,  as  are  equally  the 
Bostryx  and  the  Drepanium,  that  the  four  spe- 
cies may  as  well  be  reduced  to  two.  As  these 
severally  include  the  scorpioid  and  the  helicoid 
uniparous  cymes  of  modern  anthotaxy,  these 
terms  may  be  retained  to  designate  them.  Or, 
if  other  terms  in  use  be  preferred  to  scorpioid 
and  helicoid,  the  form  with  two-ranked  flowers 
may  be  denominated  Cincinnal,  that  with  single- 
ranked  Bostrychoidd.  But  in  neither  type  is  the  rhachis  always  coiled 
up,  although  commonly  more  or  less  so  in  the  undeveloped  state. 

While  these  forms  generally  imitate  racemes  or  spikes,  it  will  be  noted 
that  Fig.  300  specially  imitates  a  corymb  in  form  and  in  seeming  acropetal 
or  centripetal  evolution.  And  when,  as  in  this  figure,  the  bracts  are  all 
absent,  no  obvious  external  difference  remains. 

FIG.  305".  Ground  plan  (from  Eichler's  Bluthendiagramme,  i.  38)  of  the  scorpioid 
inflorescence  of  Tradescantia  erecta,  between  bract  (/?)  below  and  axis  (a)  above: 
I.,  II.,  III.,  &c.,  the  successive  flowers:  «'  is  the  bractlet  of  the  first  and  bract  of  the 
second  flower,  and  so  the  others  in  succession  up  to  v"  and  a  small  undeveloped  one 
beyond.  The  figure  1  affixed  to  each  flower  indicates  the  first  floral  leaf. 


158  ANTHOTAXY,   OK   INFLORESCENCE. 

or  a  pedicel  or  peduncle  above  and  out  of  connection  with  the 
leaf  which  should  subtend  and  accompany  it.1 

287.  Mixed  Inflorescence  is  not  uncommon.  This  name  is 
given  to  clusters  or  ramifications  in  which  the  two  types  are  con- 
joined. Being  heteromorphous,  they  are  almost  necessarily  com- 
pound, the  two  types  belonging  to  different  orders  of  ramification. 
But  under  it  may  be  included  cases  of  comparatively  simple 
inflorescence,  at  least  in  the  beginning,  some  of  which  nearty 
fuse  the  two  types  into  one.  In  the  Teasel  (Dipsacus) ,  an  appar- 
ently simple  head  or  short  spike  comes  first  into  flower  at  the 
middle,  from  which  the  flowering  proceeds  regularly  to  the  base. 
Had  it  begun  at  the  top,  it  would  answer  to  Fig.  281,  which, 
blossoming  from  above  downward  by  simple  uniflorous  lateral 
axes  along  a  monopodial  primary  axis,  is  a  simple  racemiform 
cyme,  while  it  may  also  be  called  a  reversed  or  determinate  raceme. 
Something  of  this  sort  may  be  seen  in  certain  species  of  Cam- 
panula, with  virgate  inflorescence,  the  terminal  blossom  earliest, 
the  others  following  irregularly,  or  partly  downward  and  partly 
upward.  In  C.  rapunculoides,  when  rather  depauperate  and 
£he  inflorescence  simple,  the  evolution  is  that  of  a  true  raceme, 
except  that  a  flower  at  length  terminates  the  axis  and  develops 
earlier  than  the  upper  half  of  the  raceme.  In  Liatris  spicata 
and  its  near  relatives,  the  heads,  on  the  virgate  general  axis, 
come  into  flower  in  an  almost  regular  descending  order,  or  are 
reversely  spicate.  If  in  Fig.  281  the  lower  pedicels  were  prolonged 
to  the  level  of  the  upper,  a  simple  corymbiform  cyme  would  be 
seen,  with  simple  centrifugal  evolution,  that  is,  regularly  from 
the  centre  to  the  margin  ;  this  is  the  counterpart  of  the  rhipidium 
or  fan-shaped  cyme,  of  Fig.  300,  in  which  the  evolution  of  the 
blossoms  is  as  regularly  centripetal.  The  explanation  of  the 
paradox  is  not  far  to  seek. 

1  The  position  of  a  pedicel  at  the  side  of  a  bract  in  false  racemes  is  ex- 
plained in  the  foregoing  note.  It  may  occur  in  true  racemose  inflorescence 
by  the  reduction  of  sessile  secondary  racemes  down  to  an  umbel  of  two 
flowers,  transverse  to  the  bract  (as  in  many  species  of  Desmodium),  and 
thus  seemingly  lateral  to  it,  or  to  a  single  flower  on  the  right  or  left  of  it. 
The  coalescence  of  a  pedicel  to  the  axis  for  a  considerable  height  above  the 
subtending  bract  in  a  simple  inflorescence,  or  above  the  last  leaf  in  a 
sympodial  one  (concaulescence  of  Schimper),  is  common.  So  likewise  bracts 
or  leaves  may  be  for  a  good  distance  adnate  to  sympodial  shoots,  whether 
peduncles  or  leafy  flowerless  branches.  This  (named  recaulescence  by 
Schimper)  is  of  most  frequent  occurrence  in  Solanaceae  (in  Datura,  Atropa, 
most  species  of  Solanum,  &c.),  and  is  the  explanation  of  their  so-called 
gemmate  leaves,  where  a  large  leaf  (really  belonging  lower  down)  has  a 
small  leaf  by  the  side  of  it.  See  Wydler  in  Bot.  Zeit.  ii.  689,  &c.,  Sendtner 
in  Fl.  Bras.  x.  183,  and  Eichler,  Bluthend.  i.  199. 


THE   CYMOSE  TYPE.  159 

288.  Compound  mixed  inflorescence  is  very  various  and  com- 
mon ;  but  the  combinations  have  hardly  called  for  special  terms, 
being  usually  disposed  of  by  a  separate  mention  of  the  general 
and  of  the  partial  anthotaxy,  or  that  of  the  main  axis  of  inflo- 
rescence and  that  of  its  ensuing  ramification.1  In  Composite, 
for  instance,  the  flowers  are  always  in  true  heads,  of  centripetal 
evolution.  The  heads  terminate  main  stems  as  well  as  lateral 
branches,  so  that  they  are  centrifugally  or  cymosely  disposed. 
The  reverse  occurs  in  all  Labiatae  and  most  Scrophulariaceae, 
where  the  flowers,  when  clustered,  are  in  cymes,  but  these  cymes 
are  from  axils,  and  develop  in  centripetal  order.  It  is  this 
arrangement  which  mainly  characterizes  the 

THYRSUS.  A  compound  inflorescence  of  more  or  less  elongated 
shape,  with  the  primary  ramification  centripetal  or  botryose,  the 
secondary  or  the  ultimate  centrifugal  or  cymose.  To  the  defini- 
tion is  generally  added,  that  the  middle  primary  branches  are 
longer  than  the  upper  and  lower,  rendering  the  whole  cluster 
narrower  at  top  and  bottom,  and  sometimes  that  it  is  compact : 
but  these  particulars  belong  only  to  typical  examples,  such  as 
the  inflorescence  of  Lilac  and  Horsechestnut.  In  the  former,  the 
thyrsus  is  usually  compound.  A  loose  thyrsus  is  a 

MIXED  PANICLE.  It  is  seldom  that  a  repeatedly  branching 
inflorescence  of  the  paniculate  mode  is  of  one  type  in  all  its 
successive  ramifications.  Either  the  primarily  centripetal  will 
become  centrifugal  in  the  ultimate  divisions,  or  the  primarily 
centrifugal  will  by  suppression  soon  run  into  false  botryose 
forms,  into  apparent  racemose  or  spicate  subdivisions.  So  that 
the  name  Panicle  in  terminology  is  generally  applied  to  all  such 
mixed  compound  inflorescence,  as  well  as  to  the  homogeneously 
botryose.  (278.) 

VERTICILLASTER  is  a  name  given  to  a  pair  of  opposite  and 
sessile  or  somewhat  sessile  cymes  of  a  thyrsus  or  thyrsiform 
inflorescence,  which,  when  full,  seem  to  make  a  kind  of  verticil  or 
whorl  around  the  stem,  as  in  very  many  Labiatae.  The  name 
was  originally  given  to  each  one  of  the  pair  of  cymes  ;  but  it  is 
better  and  more  commonly  used  to  denote  the  whole  glomerule 
or  false  whorl  produced  by  the  seeming  confluence  of  the  two 
clusters  into  one  which  surrounds  the  stem. 


1  Guillaud  (in  his  memoir  on  Inflorescence,  published  in  Bull.  Soc.  Bot. 
France,  iv.)  proposes  to  designate  as  Cymo-Botryes  the  mixed  inflorescence 
composed  of  cymes  developed  in  botryose  order,  i.  e.  the  thyrsus ;  and 
Botry-Cymes,  the  reverse  case  of  racemes,  &c.,  cymosely  aggregated.  For 
the  former,  the  old  name  thyrsus  serves  appropriately  and  well. 


160  ANTHOTAXY,  OR   INFLORESCENCE. 

289.  The  Relations  of  Bract,  Bractlet,  and  Flower  should  here 
be  noticed,  although  the  subject  in  part  belongs  rather  to  the 
section  on  Floral  Symmetry.  (315.) 

290.  Anterior   and   Posterior,   otherwise   called   Inferior  and 
Superior,  and  therefore  Lower  and  Upper,1  are  primary  relations 
of  position  of  an  axillary  flower  with  respect  to  subtending  bract 
and  the  axis  to  which  the  bract  pertains.     The  flower  is  placed 
between  the  two.     The  portion  of  the  flower  which  faces  the 
subtending  bract  is  the  anterior,  likewise  called  inferior  or  lower. 

The  opposite  portion  which  faces  the  axis  of 
inflorescence  is  the  posterior,  or  superior,  or 
upper.  The  right  and  left  sides  are  lateral. 
(Fig.  304,  306.)  These  relations  do  not 

)fc/appear  in  a  solitary  flower  terminating  a 
simple  stem  ;  but  when  such  an  axis  produces 
axillary  branches  with  a  terminal  flower,  the 
relation  of  this  flower  to  the  preceding  axis 
and  its  leaf  is  manifest,  just  as  in  indetermi- 
nate inflorescence. 

291.  Median  and  Transverse.     The  position  of  parts  which  lie 
in  antero-posterior  line,  or  between  bract  and  axis,  is  median. 
Thus,  in  Fig.  304  and  305,  the  parts  are  all  in  the  median  plane  : 
in  Fig.  306,  the  bractlets,  b',  b',  are  lateral  or  collateral,  or  (being 
in  the  opposite  plane)  transverse. 

292.  Position  of  Bractlets.  The  rule  has  already  been  laid 
down  (285)  that  the  first  leaf  of  an  alternate-leaved  secondary 
axis  is  in  Monoctyledons  usually  median  and  posterior,  that  is, 
farthest  away  from  the  subtending  leaf  (as  in  Fig.  304,  305)  ; 
in  Dicotyledons,  lateral  or  transverse.     When  these  secondary 
axes  are  one-flowered  peduncles  or  pedicels,  the  leaf  or  leaves 
(if  any)  they  bear  are  bractlets.2     Commonly  there  is  only  this 


1  Not  (with  propriety,  although  the  terms  have  been  so  used)  exterior  or 
outer  for  the  anterior,  and  interior  or  inner  for  the  posterior  position.     These 
terms  should  be  reserved  for  the  relative  position  on  the  axis  of  successive 
circles  or  parts  of  circles,  spirals,  &c.     Covering  or  overlapping  parts  are 
exterior  or  outer  in  respect  to  those  overlapped. 

2  Latin  Bracteolce :  not  that  they  are  small  bracts,  but  bracts  of  an  ulti- 
mate axis.    In  axillary  inflorescence,  the  distinction  between  bractlet  and 
bract  is  obvious :  in  case  of  a  solitary  terminal  flower,  there  is  no  ground  of 
difference :  in  terminal  or  cymose  inflorescence,  the  difference  is  arbitrary  ; 
but  we  may  restrict  the  term  bractlet  to  the  last  bract  or  pair. 

German  botanists  mostly  distinguish  between  bracts,  as  a  leaf  subtending 
a  flower  or  cluster,  and  bractlets,  by  terming  the  former  a  Deckblatt,  and  the 

FIG.  306.    Diagram  (cross  section)  of  papilionaceous  flower  and  its  relation  to  axis 
(a),  bract  (b),  and  bractlets  (6',  V). 


THE  CYMOSE  TYPE.  161 

posterior  one  to  a  simple  axis  in  Monocotyledons,  and  two 
transverse  ones  in  Dicotyledons,  i.  e.  one  to  the  right  and  the 
other  to  the  left  of  the  subtending  bract,  Fig.  306,  V  b'.  When 
the  latter  form  a  pair,  they  are  perhaps  always  truly  transverse ; 
when  alternate,  they  stand  more  or  less  on  the  opposite  sides  and 
transverse.  When  more  than  one  in  Monocotyledons,  they  may 
become  either  median  or  transverse,  or  even  intermediate.  The 
relation  of  bractlets  or  bract,  that  is,  of  the  last  leaves  of  inflo- 
rescence, to  the  first  of  the  blossom,  might  be  considered  either 
under  Phyllotaxy  or  under  Floral  Symmetry.  In  general,  it 
may  be  noted  that  successive  members  stand  over  the  widest 
intervals  ; 1  in  other  words,  that  the  first  leaf  of  the  flower  is  as 
far  away  as  may  be  from  the  highest  bractlet.  For  instance, 
when  there  is  a  single  and  posterior  bractlet,  as  is  common  in 
Monocotyledons,  the  first  leaf  of  the  flower  is  anterior,  the 
next  two  right  and  left  at  120°.  When  there  is  a  single  and 
lateral  bractlet  and  five  leaves  in  the  first  circle  of  the  flower 
(which  occurs  only  in  Dicotyledons) ,  the  first  leaf  of  this  circle 
is  either  exactly  on  the  opposite  side  from  the  bract,  or  at  a 
divergence  of  two  fifths,  the  latter  falling  into  the  continuous 
spiral.  When  with  a  pair  of  bractlets,  right  and  left,  the  first 
flower-leaf  is  at  £  divergence  from  one  (the  uppermost)  of  them 
when  the  circle  is  of  three,  or  at  £  when  of  five  members,  or  near 
it ;  but  with  many  exceptions.2 

A  tabular  view  of  the  kinds  of  inflorescence  and  their  termi- 
nology, serving  as  a  key,  may  aid  the  student.8 


latter,  being  the  leaves  which  the  new  axis  first  bears,  Vbrblatter,  which  is 
also  the  name  they  apply  to  primordial  leaves  in  germination. 

1  In  accordance  with  Hofmeister's  law  ;  but  (as  Eichler  remarks)  not  to 
be  explained  on  his  mechanical  principle  of  production  in  this  place  because 
of  the  greater  room :  for  the  position  of  the  first  member  of  an  axillary 
flower  is  mostly  the  same  as  regards  the  subtending  bract  when  the  bractlets 
are  wanting. 

2  When  bractlets  are  wanting,  the  leaves  of  the  first  floral  circle  if  two 
are  right  and  left ;  if  three,  two  lateral-posterior  and  one  anterior ;  when 
five,  the  odd  one  commonly  in  the  median  line,  either  anterior  or  posterior. 


8  INFLORESCENCE  is  either  PURE,  all  of  one  type,  or  MIXED,  of  the  two  types 

combined.  The  Types  are : 

I.  Main  axis  not  arrested  and  terminated  by  a  flower.    Indeterminate,  Indefi- 
nite, Acropetal  or  Ascending,  Centripetal,  or  BOTRYOSE. 

II.  Main  and  lateral  axes  arrested  and  terminated  by  a  flower.    Determinate, 
Definite,  Descending,  Centrifugal,  or  CYMOSE. 


162  ANTHOTAXY,  OB  INFLORESCENCE. 


L  BOTRYOSE  TYPE. 

1.  Simple,  with  lateral  axes  unbranched  and  terminated  by  a  single  flower,  and 

Flowers  on  pedicels, 

Of  somewhat  equal  length  on  a  comparatively  elongated  axis,    RACEME. 

The  lower  ones  longer  than  the  upper,  and  main  axis  short,  .    CORYMB. 

Of  nearly  equal  length  on  an  undeveloped  main  axis,      .    .    UMBEL. 

Flowers  sessile  on  a  very  short  main  axis, HEAD. 

Flowers  sessile  on  a  comparatively  elongated  main  axis, .    .     .    SPIKE. 

A  fleshy  spike  or  head  is  a SPADIX. 

A  scaly-bracted  spike  is  an AMENT  or 

CATKIN. 

2.  Compound,  with  lateral  axes  branched  once  or  more,  bearing  clusters  instead  of 

single  flowers. 

Irregularly  racemosely  or  corymbosely  compound,  .     .  PANICLE. 
Homogeneously  and  regularly  compound,  as 

Racemes  in  a  raceme, COMPOUND  RACEME. 

Corymbs  corymbose, COMPOUND  CORYMB. 

Umbels  in  an  umbel, COMPOUND  UMBEL. 

Spikes  spicate, COMPOUND  SPIKE. 

Homogeneously  compound,  the  secondary  ramification 

unlike  the  primary,  as  Heads  racemose,  Umbels  spiked, 

Spikes  panicled,&c. 

II.  CYMOSE  TYPE. 

1.  Simple,  with  terminal  axis  of  each  generation  one-flowered. 

Monopodial,  the  axis  of  each  generation  evidently  re- 
solved into  branches, TRUE  CYME. 

These  more  than  two, Pleiochasium  or  MULTIPAROUS  CYME. 

These  only  two,      .    .    .  Dichasium,  Dichotomous  or  BIPAROUS  CYME. 
Sympodial,  the  apparently  simple  axis  continued  by 
a  succession  of  new  axes  standing  end  to  end, 
Monochasium,  False  Raceme  or  Spike,  Botryose  or  UNIPAROUS  CYMB. 
Flowers  one-ranked  on  one  side  of  rhachis,    HELICOID  UNIPAROUS  CYME. 
Flowers  two-ranked  on  one  side  of  rhachis,   SCORPIOID  UNIPAROUS  CYME. 

2.  Compound,  with  terminal  axes  for  one  or  more  earlier 

generations  bearing  a  cyme  instead  of  a  single 

flower, Various  sorts  of  COMPOUND  CYME. 

III.  MIXED  INFLORESCENCE. 

1.  Anomalous  simple,  with  unbranched  one-flowered  lateral 

axes  (287),  such  as    ...  PARTLY  REVERSED  SPIKES  OR  RACEMES 

2.  Compound,   of  various  combinations,  of  which  there  are 

names  for  the  subjoined :  — 
Primary  inflorescence  botryose,  with  axis  elongating; 

secondary  cymose, THYRSUS. 

Pair  of  such  opposite  cymes  seemingly  confluent  round 

the  main  axis, VERTICILLASTER. 

Panicle  with  some  of  the  ramifications  cymose,    .    .    .  MIXED  PANICLE. 


THE  FLOWEB.  163 


CHAPTER  VI. 

THE  FLOWEB. 

SECTION  I.     ITS  NATURE,  PARTS,  AND  METAMORPHT. 

293.  FLOWER- BUDS  are  homologous  with  (morphologically  an- 
swering to)  leaf-buds,  and  they  occupy  the  same  positions.  (266.) 
A  FLOWER  is  a  simple  axis  or  a  terminal  portion  of  one,  in  a 
phaenogamous  plant,  with  its  leaves  developed  in  special  forms, 
and  subservient  to  sexual  reproduction  instead  of  vegetation. 

294.  In  passing  from  vegetation  to  reproduction,  it  is  not 
always  easy  to  determine  exactly  where  the  flower  begins.     The 
same  axis  which  bears  a  flower  or  floral  organs  at  summit  bears 
vegetative  leaves  or  foliage  below.     Or  when  it  does  not,  as 
when  an  axillary  flower-stalk  or  pedicel  is  bractless,  the  change 
to  actual  organs  of  reproduction  is  seldom  abrupt.     Usually 
there  are  floral  envelopes,  within  and  under  the  protection  of 
which  in  the  bud  the  essential  organs  of  the  flower  are  formed. 
Some  or  all  of  these  protecting  parts,  in  very  many  flowers,  are 
either  obvious  leaves  or  sufficiently  foliaceous  to  suggest  their 
leafy  nature ;  and  even  when  the  texture  is  delicate,  and  other 
colors  take  the  place  of  the  sober  green  of  vegetation,  they  are 
still  popularly  said  to  be  the  leaves  of  the  blossom.     These  pro- 
tecting and  often  showy  parts,  though  not  themselves  directly 
subservient  to  reproduction,  have  always  been  accounted  as 
parts  of  the  flower.1    Between  the  lowest  or  outermost  of  these 
and  the  bractlets  and  bracts  there  are  various  and  sometimes 
complete  gradations.     The   axis  itself  occasionally  undergoes 
changes  in  such  a  way  as  to  render  the  determination  of  the 
actual  beginning  of  the  flower  somewhat  arbitrary.     Moreover, 
the  flower  itself  is  extremely  various  in  different  plants,  in  some 
consisting  of  a  great  number  of  pieces,  in  others  of  few  or  only 
one ;   in  some  the  constituent  pieces  are  separate,  in  others 
combined.     The  flower  is  best  understood,  therefore,  by  taking 
some  particular  specimen  or  class  of  flowers  as  a  representative 

1  Indeed,  the  colored  leaves,  or  envelopes  in  whatever  form,  essentially 
were  the  flower  in  most  of  the  ante-Linnaean  definitions  (that  of  Ludwig 
excepted),  as  they  are  still  mainly  so  in  popular  apprehension. 


164 


THE  FLOWER. 


or  pattern,  and  especially  some  one  which  is  both  complete  and 
morphologically  simple. 

295.  Such  a  flower  consists  of  two  kinds  of  organs,  viz.  the 
Protecting  Organs,  leaves  of  the  blossom,  or  floral  envelopes, 
which,  when  of  two  sets,  are  CALYX  and  COROLLA  ;  and  the 
Essential  Reproductive  Organs,  which  co-operate  in  the  production 
of  seed,  the  STAMENS  and  PISTILS. 

296.  Floral  Envelopes,  Perianth,  or  Perigone,  the  floral  leaves 
or  coverings.     The  former  is  a  proper  English  designation  of 
these  parts,   taken  collectively.      But  in  descriptive  botany, 
where  a  single  word  is  preferable,  sometimes  the  name  perianth 
(Lat.  perianthium) ,  sometimes  that  of  perigone  (or  perigonium) , 
is  used.     Perianthium,1  a  Linnsean  term,  has  been  objected  to, 
because  it  etymologically  denotes  something  around  the  flower ; 
but  it  seems  not  inappropriate  for  the  envelopes  which  surround 

the  essential  part  of  the  flower.  Perigonium, 
a  later  term,  has  the  advantage  of  meaning 
something  around  the  reproductive  organs, 
which  is  precisely  what  it  is.  Neither  name 
is  much  used,  except  where  the  perianth  or 
perigone  is  simple  or  in  one  set  (when  it  is 
almost  always  cah^) ,  or  where  it  is  of  two 
circles  having  the  gen- 
eral appearance  of  one 
and  needing  descrip- 
tive treatment  as  such, 
as  in  the  petaloideous 
Monocotyledons.  It  is 
also  used  where  the 
morphology  is  ambigu- 
ous. Generally,  the  floral  envelopes  are  treated  distinctively  as 
calyx  and  corolla,  one  or  the  other  of  which  (mostly  the  corolla) 
may  be  wanting. 

297.  The  Calyx  is  the  outer  set  of  floral  envelopes.    That  is 
its  only  definition.    Commonly  it  is  more  herbaceous  or  foliaceous 
than  the  corolla,  and  more  persistent,  yet  sometimes,  as  in  the 
Poppy  family,  it  is  the  more  deciduous  of  the  two.     Not  rarely  it 

1  Linnaeus  (and  about  the  same  time  Ludwig)  used  it  in  the  sense  of  a 
proper  calyx,  yet  with  some  vagueness.  Mirbel  and  Brown  established  it 
in  the  sense  of  the  collective  floral  covering.  DeCandolle  revived  Ehrhart's 

PIG.  307.  The  complete  flower  of  a  Crassula.  308.  Diagram  of  its  cross-section  in 
the  bud,  showing  the  relative  position  of  its  parts.  The  five  pieces  of  the  exterior 
circle  are  sections  of  the  sepals;  the  next,  of  the  petals;  the  third,  of  the  stamens 
through  their  anthers;  the  innermost,  of  the  five  pistils. 

FIG.  309.  A  sepal;  310,  a  petal;  311,  a  stamen;  and  312,  a  pistil  from  the  flower 
represented  in  Fig.  307. 


ITS   NATURE   AND    PARTS.  165 

is  as  highly  colored.  A  name  being  wanted  for  the  individual 
leaves  which  make  up  the  calyx,  analogous  to  that  for  corolla- 
leaves,  DeCandolle  adopted  Necker's  coinage  of  the  word  sepal. 
Calyx-leaves  are  SEPALS. 

298 .  The  Corolla  is  the  inner  set  of  floral  envelopes,  usually  (but 
not  always)  of  delicate  texture  and  other  than  green  color,  form- 
ing therefore  the  most  showy  part  of  the  blossom.     Its  several 
leaves  are  the  PETALS.1 

299.  The  floral  envelopes  are  for  the  protection  of  the  organs 
within,  in  the  bud  or  sometimes  afterward.     Also,  some  of  them, 
by  their  bright  colors,  their  fragrance,  and  their  saccharine  or 
other  secretions,  serve  for  allurement  of  insects  to  the  blossom,  to 
mutual  advantage.  (504.)     This  furnishes  a  reason  for  neutral 
flowers,   those   devoid   of  essential    organs,    which    sometimes 
occur  along  with  less  conspicuous  perfect  ones.     "The leaves  of 
the  flower  "  are  therefore  indirectly  subservient  to  reproduction. 

300.  The  essential  organs,  being  commonly  plural 
in  number,  sometimes  need  a  collective  name.  Where- 
fore, the  aggregate  stamens  of  a  flower  have  been 
called  the  ANDRCECIUM  ;  the  pistils,  the  GYN<ECiUM.2 

301.  The  Stamens8    are  the  male  or  fertilizing 
organs  of  a  flower.     A  complete  stamen  (Fig.  311, 
313)  consists  of  FILAMENT  (/),  the  stalk  or  support, 
and  ANTHER  (a),  a  double  sac  or  body  of  two  cells, 

side  by  side,  filled  with  a  powdery  substance,  POLLEN,  which  is 
at  length  discharged,  usually  through  a  slit  or  cleft  of  each  cell. 

well-formed  name  of  perigonium,  and  in  the  sense  here  given.  But  later  (in 
the  Organographie)  he  proposed  to  restrict  it  to  cases  in  which  the  part  is 
of  ambiguous  nature,  as  in  Monocotyledons.  The  earlier  definition  is  no 
doubt  the  proper  one ;  but  the  occasions  for  using  the  term  in  descriptive 
botany  are  mainly  where  the  nature  may  seem  to  be  ambiguous  or  con- 
fused, or  where,  from  the  union  or  close  similarity  of  outer  and  inner  circles, 
it  is  most  convenient  to  treat  the  parts  as  forming  one  organ. 

1  Fabius  Columna,  at  the  close  of  the  sixteenth  century,  appears  to  have 
introduced  this  term,  or,  as  Tournefort  declares,  "primus  omnium  quod 
sciam  Petali  vocem  proprie  usurpavit,  ut  folia  florum  a  foliis  proprie  dictis 
distingueret." 

2  The  male  household  and  the  female  household  respectively,  terms  in- 
troduced by  Keeper  (Linnaea,  i.  437),  in  the  form  of  andrceceum  &n(igyn(Eceum ; 
but  the   diphthong  in  the   latter  should   also   be  a.      The    orthography 
andrcecium  and  gynoecium  (early  adopted  by  Bentham,  in  Labiatarum  Gen.  et 
Spec. )  is  conformable  to  the  Linnaean  Monaxia,  Dicecia,  &c. 

8  The  name  (from  the  Greek  and  Latin  name  of  the  warp  of  the  ancient 
upright  loom,  and  thence  used  in  the  sense  of  threads)  was  applied,  down 
to  Tournefort  and  later,  to  the  filaments;  and  the  anthers  were  termed 

FIG.  313.  Stamen,  composed  of/,  filament,  and  a,  anther,  with  cells  opening 
laterally  and  discharging  pollen. 


166 


THE  FLOWER. 


302.  The  Pistils,  one  or  more  to  the  flower,  are  the  female  or 
seed-bearing  organs.1     A  complete  pistil  is  distinguished  into 
three     parts :     the     OVARY     (Lat. 
Ovarium,  Fig.  314,  a,  shown  in  verti- 
cal section,  and  Fig.  3 15,  by  Linnaeus 
named  Germeri),  the  hollow  portion 
at    the    base    which    contains    the 
OVULES,  or  bodies  destined  to  be- 
come seeds  ;  the  STYLE  (&),  or  colum- 
nar prolongation  of  the  apex  of  the 
ovary ;  and  the  STIGMA  (c),  a  portion 
of  the  surface  of  the  style  denuded  of  epidermis, 
sometimes   a  mere   point  or  a   small  knob   at  the 
apex  of  the  style,  but  often  forming  a  single  or 
double  line  running  down  a  part  of  its  inner  face, 
and   assuming  a  great  diversity  of  appearance  in 
different  plants.     The  ovary  and  the  stigma  are  the 
essential  parts.     The  style  (as  also  the  filament  of  a 
stamen)  may  be  altogether  wanting. 


apices.  It  came  in  time  to  be  used  as  now  for  the  whole  organ ;  but  Lud- 
wig  (Inst.  Reg.  Veg.),  in  1742,  apparently  first  so  defined  it,  and  introduced 
the  term  Anther  for  the  Apex  of  Ray,  or  Theca  of  Grew. 

1  Following  Linnaeus,  this  term  is  here  freely  used  in  the  plural,  and  for 
each  actual  separate  member  of  the  gynoecium,  each  organ  which  has  an 
ovary,  stigma,  and  commonly  a  style.  Tournefort,  who  appears  to  have 
introduced  the  word,  employed  it  hi  the  sense  of  gynoecium.  Many  authors 
define  it  thus,  and  then  practically  eliminate  from  botany  this,  one  of  the 
oldest  of  its  terms,  and  one  by  no  means  superfluous.  The  typical  pistittum 
of  Tournefort  is  that  of  the  Crown  Imperial  (Inst.  i.  69,  &  tab.  1)  and  the 
name  is  from  the  likeness  to  a  pestle  in  a  mortar.  As  it  soon  became  im- 
possible to  apply  the  same  name  to  the  pistil  of  a  Fritillaria  or  of  a  Plum, 
the  cluster  of  such  organs  in  Caltha,  and  the  capitate  cluster  and  receptacle 
of  such  organs  in  a  Ranunculus  or  Anemone,  Linnaeus,  and  Ludwig  before 
him,  took  the  idea  of  Tournefort's  name,  and  used  it  accordingly. 

"  Pistittum  est  pars  interior  et  media  floris,  quae  ex  ovario  et  stylo  com- 
ponitur.  .  .  .  Ovarium  est  pars  pistilli  inferior,  quae  f uturi  fructus  delinea- 
tionem  sistit.  .  .  .  Stylus  est  pars  pistilli  ex  ovario  centre  producta.  .  .  . 
Summitas  styli  vel  ejus  partium  Stigma  dicitur."  Ludwig,  Inst.  Reg.  Veg. 
41-43,  1742.  Without  mentioning  the  plural,  the  pistil  is  thus  defined  in  a 
way  which  necessitates  its  use.  Linnaeus  (in  Phil.  Bot.)  first  defines  Stamen 
and  Pistillum  in  the  singular  number,  enumerating  the  three  parts  of  the 
latter,  and  afterwards  (p.  67)  declares  that  "Pistilla  differunt  quoad 

PIG.  314.  Vertical  section  of  a  pistil,  showing  the  interior  of  its  ovary,  a,  to  one 
side  of  which  are  attached  numerous  ovules,  d :  above  is  the  style,  b,  tipped  by  the 
stigma,  c. 

FIG.  315.  A  pistil  of  Crassula,  like  that  of  Fig.  312,  but  more  magnified,  and  cut 
across  through  the  ovary,  to  show  its  cell,  and  the  ovules  it  contains;  also  pulled  open 
below  at  the  suture.  At  the  summit  of  the  style  is  seen  a  somewhat  papillose  portion, 
destitute  of  epidermis,  extending  a  little  way  down  the  inner  face :  this  is  the  stigma. 


ITS   NATURE  AND   PARTS. 


167 


303.  The  Torus,   or  Receptacle  of  the  flower,   also  named 
THALAMUS,1  is  the  axis  which  bears  all  the  other  parts,  that 
upon    which    they   are    all  <*     c 
(mediately  or  immediately) 

inserted.  These  are  all  ho- 
mologous with  leaves.  This 
is  extremity  of  stem,  or 
floral  axis,  out  of  which  the 
organs  described  grow,  in 
succession,  like  leaves  on 
the  stem ;  the  calyx  from 
the  very  base,  the  petals 
next  within  or  above  the  calyx,  then  the  stamens,  finally  the 
pistils,  which,  whether  several  or  only  one,  terminate  or  seem  to 
terminate  the  axis.  (Fig.  316.) 

304.  Metamorphosis.      If  flower-buds   are   homologous   with 
leaf-buds,  and  the  parts  of  the  flower  therefore  answer  to  leaves 
modified  to  special  functions  (293),  then  the  kind  of  flower  here 
employed  in  explaining  and  naming  these  parts   is  a  proper 
pattern  blossom.      For   the    organs    are   all   separate   pieces, 
arranged  on  the  receptacle  as  leaves  are  on  the  stem,  the  outer- 
most manifestly  leaf- like,  the  next  equally  so  in  shape,  though 
not  in  color,  the  stamens  indeed  have  no  such  outward  resem- 
blance,  but  the   ripe   pistils  open  down   the  inner  angle  and 
flatten  out  into  a  leaf- like  form.     The  adopted  theory  supposes 
that  stamens  and  pistils,  as  well  as  sepals  and  petals,  arehomolo- 

numerum,"  etc.,  and  so  elsewhere,  besides  founding  his  orders  on  the  num- 
ber of  pistils.  Among  even  French  authors,  Mirbel  (1815)  writes,  "Le 
nombre  des  pistils  n'est  pas  le  meme  dans  toutes  les  especes,"  &c.  Moquin- 
Tandon  freely  refers  to  pistils  in  the  plural,  and  Aug.  St.  Hilaire  takes 
wholly  the  view  here  adopted,  distinguishing  the  solitary  pistil  into  simple 
and  compound.  DeCandolle,  in  Theorie  £lementaire,  third  edition,  writes, 
"  Chaque  carpel  est  un  petit  tout,  un  pistil  entier,  compose'  d'un  ovaire,  d'un 
style,  et  d'un  stigmate."  Of  English  authors,  no  other  need  be  cited  than 
Robert  Brown.  The  terms  in  question,  then,  are  :  — 

Gyncecium,  the  female  system  of  a  flower,  taken  as  a  whole. 

Pistil,  each  separate  member  of  the  gynoecium;  this  either  simple  or 
compound. 

Ovary,  the  ovuliferous  portion  of  a  pistil.  Substituting  a  part  for  the 
whole,  this  term  is  often  used  when  the  whole  pistil  is  meant. 

Carpel,  or  Carpid,  or  Carpophyll,  each  pistil-leaf ;  whether  distinct  as  in 
simple  or  apocarpous  pistils,  or  in  combination  of  two  or  more  to  form  a  com- 
pound or  syncarpous  pistil. 

1  By  Tournefort,  and  adopted  by  Ludwig.  Receptaculum  jlaris,  Linnaeus. 
Thorns,  Salisbury.  Torus  (the  proper  form),  DeCandolle. 

FIG.  316.  Parts  of  the  flower  of  a  Stonecrop,  Sedum  ternatum,  two  of  each  sort, 
and  the  receptacle,  displayed:  a,  sepal;  b,  petal;  c,  stamen;  d,  pistil. 


168  THE   FLOWER. 

gous  with  leaves ;  that  the  sepals  are  comparatively  little,  the 
petals  more,  and  the  reproductive  organs  much  modified  from  the 
type,  that  is  from  the  leaf  of  vegetation.  This  is  simply  what 
is  meant  by  the  proposition  that  all  these  organs  are  transformed 
or  metamorphosed  leaves.  What  would  have  been  leaves,  if 
the  development  had  gone  on  as  a  vegetative  branch,  have  in 
the  blossom  developed  in  other  forms,  adapted  to  other  func- 
tions. Linnaeus  expressed  this  idea,  along  with  other  more 
speculative  conceptions,  dimly  apprehended,  by  the  phrase  Vege- 
table Metamorphosis.  Not  long  afterwards,  this  fecund  idea 
of  a  common  type,  the  leaf,  of  which  the  parts  of  the  flower, 
&c.,  were  regarded  as  modifications,  was  more  clearly  and  differ- 
ently developed  by  a  philosophical  physiologist,  Caspar  Frederic 
Wolff.  Thirty  years  later,  it  was  again  and  wholly  independ- 
ently developed  by  Goethe,  in  a  long-neglected  but  now  well- 
known  essay,  on  the  Metamorphosis  of  Plants.  Twenty- three 
years  afterwards,  similar  ideas  were  again  independently  pro- 
pounded by  DeCandolle,  from  a  different  theoretical  point  of 
view ;  and  finally  the  investigation  of  phyllotaxy  has  completed 
the  evidence  of  the  morphological  unity  of  foliaceous  and  floral 
organs.1 

1  The  contribution  of  Linnaeus  is  on  p.  301  of  the  Philosophia  Botanica, 
1761 ;  and  all  that  is  pertinent  is  in  the  following  propositions  :  — 

Plumulam  seminis  saepius  terminat  aut  flos  aut  gemma. 

Principium  florum  et  f  oliorum  idem  est. 

Principium  gemmarum  et  f  oliorum  idem  est. 

Gemma  constat  f  oliorum  rudimentis. 

Perianthium  sit  ex  connatis  foliorum  rudimentis. 

His  dissertation,  Prolepsis  Plantarum,  in  Amoen.  Acad.  vi.  (1760),  added 
nothing  but  obscure  speculations  to  the  former  comparatively  clear 
statements. 

Kaspar  Friedrich  Wolff's  contribution  is  in  his  Theoria  Generationis, 
mainly  concerning  animals,  published  in  1759,  and  an  enlarged  and  amended 
edition  in  1774.  He  first  clearly  conceives  the  plant  as  formed  of  two  ele- 
ments, stem  and  leaf,  but  develops  only  the  morphology  of  the  latter,  and 
under  the  hypothesis  that  leaves  of  vegetation  become  bud-scales  or  floral 
organs,  as  the  case  may  be,  through  degenerescence  or  diminution  of  vege- 
tative force,  which  is  renewed  in  the  bud  or  in  the  seed. 

Johann  Wolfgang  Gothe's  Versuch  die  Metamorphose  der  Pflanzen  zu 
erklaren  was  published  in  1790,  in  86  pages.  For  the  translations  and 
reproductions,  see  Pritzel,  Thesaurus.  To  the  French  translation  by  Soret, 
with  German  text  accompanying  (Stuttgart,  1831),  and  also  to  that  of  Ch. 
Martens  (CEuv.  Hist.  Nat.  de  Goethe,  Paris,  1837),  are  joined  the  author's  inter- 
esting notes  and  anecdotes  of  later  periods,  down  to  1831.  The  degenerescence 
by  diminution  of  vegetative  force  with  renewals  by  generation,  propounded 
by  Wolff,  in  Goethe's  essay  takes  the  form  of  successive  expansion  and  con- 
traction of  organs. 

A.  P.  DeCandolle's  Theorie  Ele"mentaire  de  Botanique  appeared  in  1813, 


ITS   MBTAMORPHY.  169 

305.  It  will  be  understood  that  metamorphosis,  as  applied  to 
leaves  and  the  like,  is  a  figurative  expression,  adding  nothing  to 
our  knowledge  nor  to  clearness  of  expression,  but  rather  liable 
to  mislead.     The  substance  of  the  doctrine  is  unity  of  type.     Its 
proof  and  its  value  lie  in  the  satisfactory  explanation  of  the  facts, 
all  of  which  it  co-ordinates  readily  into  a  consistent  and  simple 
system.     As  applied  to  the  flower,  two  kinds  of  evidence  may 
be  adduced,  one  from  the  normal,  the  other  from  teratological 
conditions  of  blossoms.     The  principal  evidence  of  the  first  class 
is  that  supplied  by 

306.  Position  and  Transitions.     As  illustrated  in  the  preced- 
ing chapter,  the  flower  occupies  the  place  of  an  ordinary  bud  or 
leaf-bud.     Also  the  parts  of  the  flower  are  arranged  on  the 
receptacle  as  leaves  are  arranged  on  the  stem,  i.  e.  they  conform 
to  phyllotaxy,  as  well  in  passing  from  leaves  and  bracts  to  the 
perianth,  as  in  the  position  of  the  floral  organs  in  respect  to  each 
other.     This   is  partly  shown  in   the  preceding   chapters,  and 
is   to  be   further    illustrated.      Sepals,    petals,    stamens,   and 
pistils   are  either  in   whorls  or  in   spirals,  and  have   nothing 
in  their  arrangements  as  to  position  which  is  not  paralleled  in 
the  foliage. 

307.  The  evidence  from  transitions  has  to  be  gathered  from  a 
great  variety  of  plants.     Very  commonly  the  change  is  abrupt 
from  foliage  to  bracts,  from  bracts  to  calyx-leaves,  from  these  to 
corolla-leaves,  and  from  these  to  stamens.    But  instances  abound 
in  which  every  one  of  the  intervals  is  bridged  by  transitions  or 

a  second  edition  1819 ;  a  third  (revised  by  Alphonse  DeCandolle),  in  1844, 
is  posthumous.  The  Organographie  Vegetale,  in  which  the  morphology  of 
the  earlier  work  is  developed,  appeared  in  1827.  The  leading  idea  is  that 
of  symmetry,  of  organs  symmetrically  disposed  around  an  axis  (the 
homology  of  foliar  and  floral  organs  not  at  first  apprehended),  but  this 
symmetry  disguised  or  deranged  more  or  less  by  unions  (solderings)  of 
homogeneous  or  heterogeneous  parts,  by  irregularities  or  inequalities  of 
growth,  by  abortions,  &c. 

The  reason  why  the  organs  in  question  have  a  normal  symmetrical  dis- 
position on  the  vegetative  and  floral  axes  was  not  reached  by  DeCandolle, 
nor  was  it  perceived  that  the  arrangement  of  leaves  and  of  floral  organs  was 
identical.  All  this  was  the  contribution  of  phyllotaxy,  —  a  subject  which 
was  approached  by  Bonnet  (an  associate  of  DeCandolle's  father),  and  first 
investigated  by  the  late  Karl  Schimper  and  Alexander  Braun,  beginning 
about  the  year  1829. 

It  is  interesting  to  know  that  Wolff's  work  was  wholly  unknown  to 
Goethe  in  1790,  and  that  both  Wolff's  and  Goethe's  were  unknown  to  DeCan- 
dolle until  after  the  publication  of  the  second  edition  of  the  latter's  The'orie 
El&nentaire,  in  1819.  When  the  Organographie  appeared,  the  essay  of  Goethe 
had  come  to  light ;  and  contemporary  contributions  to  floral  morphology  by 
Petit-Thouars,  R.  Brown,  Dunal,  and  Boeper,  were  adding  their  influence. 


170  THE  FLOWER. 

intermediate  forms.     The  gradual  transition  from  ordinary  foli- 
age to  bracts  and  bractlets  is  exceedingly  common.     In  color 

and  texture  it  is  not  rare  to 
meet  with  bracts  which  vie 
-with,  or  indeed  surpass,  pet- 
als themselves  in  delicacy  and 
brightness  ;  and  in  such  cases 
they  assume  a  principal  office 
of  flower-leaves,  that  of  con- 
spicuous show  for  attraction. 
Scarlet  Sage,  Painted-Cup 
(Castilleia) ,  and  the  Poin- 
settia,  with  other  Euphorbias 
of  the  conservatories,  are  ex- 
amples of  this .  In  the  flowers 
yntd  &£j  of  Barberry,  it  is  by  a  nearly 
arbitrary  selection  that  bractlets  are  distinguished  from  sepals  ; 
in  Calycanthus,  in  many  kinds  of  Cactus,  and  in  Nelumbium, 

the    same    is    true 
as  to  bractlets,  se- 
pals, and  petals ;  in 
Water-Lily    (Nym- 
phsea,    Fig.    318), 
there  is  a  gradual 
transition  from  the 
sepals  through   the 
stamens ; 
and  most 
flowers,  se- 

1  /         pals  are  as  brightly 
\H  colored    as    petals, 

and  commonly  more 
or  less  combined 
with  them.  When 
the  perianth-leaves  are  of  only  one  set,  it  is  not  at  all  by  color  or 
texture  that  this  perianth  can  be  assigned  to  calyx  or  to  corolla. 
Normal  transitions  from  a  stamen  to  a  pistil  could  not,  in  the 
nature  of  the  case,  be  expected. 

308.  Teratological  Transitions  and  Changes.     Teratology  is 
the  study  of  monstrosities.    These  in  the  vegetable  kingdom 


FIG.  317.    Cactus-flower  (Mamillaria  caespitosa),  with  bractlets,  sepals,  and  petals 
passing  into  each  other. 

FIG.  318.    Series  exhibiting  transition  from  sepals  to  stamens  in  Nymphsea  odorata. 


m  /H  petals  to 
I  M  in  Lilies 
i  /#  lily-like  fl< 


ITS  METAMOKPHY. 


171 


often  elucidate  the  nature  of  organs.1  The  commonest  of  these 
changes  belong  to  what  was  termed  by  Goethe  retrograde  meta- 
morphosis ;  that  is,  to  reversion  from  a  higher  to  a  lower  form,  as  of 
an  organ  proper  to  the  summit  or  centre  of  the  floral  axis  into  one 
which  belongs  lower  down.2  The  most  familiar  of  all  such  cases 
is  that  of  the  so-called  double  flower,  better  named  in  Latin  flos 
plenus.  In  this,  the  essential  organs,  or  a  part  of  them,  are 
changed  into  colored  flower-  leaves  or  petals.  Most  flowers  are 
subject  to  this  change  under  long  cultivation  (witness  "  double" 
roses,  camellias,  and  buttercups)  ,  at  least  those  with  numerous 
stamens.  It  occasionally 
occurs  in  a  state  of  nature. 
The  stamens  diminish  as 
the  supernumerary  petals 
increase  in  number  ;  and 
the  various  bodies  that  may 
be  often  observed,  inter- 
mediate between  perfect 
stamens  (if  any  remain)  and 


the  outer  row  of  petals,  — 

from  imperfect  petals,  with 

a  small  lamina  tapering  into 

a   slender    stalk,   to   those 

which  bear  a  small  distorted 

lamina  on  one  side  and  a 

half-formed  anther  on  the 

other,  —  plainly  reveal  the 

nature  of  the  transformation 

that  has  taken  place.     Carried  a  step  farther,  the  pistils  likewise 

disappear,  to  be  replaced  by  a  rosette  of  petals,  as  in  fully  double 


1  The  leading  treatises  are  Moquin-Tandon's  Te'ratologie  Vegetale,  Paris, 
1841,  and  Masters,  Vegetable  Teratology,  London,  published  for  the  Ray 
Society,  1869.     An  earlier  publication  deserves  particular  mention,  viz.  the 
thesis  De  Antholysi  Prodromus,  by  Dr.  George  Engelmann,  Frankfort  on 
the  Main,  1832. 

2  To  these  abnormal  changes,  the  term  metamorphosis  is  obviously  more 
applicable ;  for  here  what  evidently  should  be  stamens,  pistils,  &c.,  on  the 
testimony  of  position  and  the  whole  economy  of  the  blossom,  actually  ap- 
pear in  the  form  of  some  other  organ  :  yet  even  here  the  change  is  only  in 
the  nisus  formativus ;  the  organ  was  not  first  formed  as  a  stamen,  and  then 
transformed  into  a  petal  or  leaf. 

FIG.  319.  A  flower  of  the  common  White  Clover  reverting  to  a  leafy  branch ;  after 
Turpin.  Calyx  with  tube  little  changed,  but  lobes  bearing  leaflets.  Pistil  stalked; 
the  ovary  open  down  the  inner  edge,  and  the  margins  of  the  pistil-leaf  bearing  leaves 
instead  of  ovules. 


172  THE   FLOWER. 

buttercups.1  In  these  the  green  hue  of  the  centre  of  the  rosette 
indicates  a  tendency  to  retrograde  a  step  farther  into  sepals,  or 
into  a  cluster  of  green  leaves.  This  takes  place  in  certain  blos- 
soms of  the  Strawberry,  the  Rose,  &c.  Such  production  of 
"  green  roses,"  and  the  like,  has  been  appropriately  called 
chlorosis,  or  by  Masters  chloranthy,  from  the  change  to  green. 
309.  A  monstrosity  of  the  blossom  of  White  Clover,  long  ago 
figured  by  Turpin  (Fig.  319),  is  such  a  case  of  foliaceous  rever- 
sion, in  which  even  the  ovules  are  implicated. 
The  imperfect  leaves  which  take  the  place  of 
the  latter  may  be  compared  with  the  leafy 
tufts  which  form  along  the  margins  of  a  leaf 
of  Bryophyllum,  by  which  the  plant  is  often 
propagated.  (Fig.  322.) 

310.  The  reversion  of  a  simple  pistil  di- 
rectly to  a  leaf  is  seen  in  the  Double-flowering 
Cherry  of  cultivation  (Fig.  320,  321),  usually 
passing  moreover,  by  prolification  of  the  re- 
ceptacle, into  a  leafy  branch. 

311.  The  reversion  of  pistils  to  stamens  is 
rarer,  but  has  been  observed  in  a  good  number 
of  instances,  in  Chives,  in  the  Horseradish, 

in  Gentians  and  Hyacinths,  and  in  some  Willows.  In  the 
latter,  the  opposite  transformation,  of  stamens  to  carpels,  is 
very  common,  and  curious  grades  between  the  two  are  met  with 
almost  every  spring.  So  also  in  the  common  Houseleek,  and 
in  perennial  Larkspurs.  Certain  apple-trees  are  known,  both 
in  the  United  States  and  Europe,  in  which,  while  the  petals  are 
changed  into  the  appearance  of  minute  green  sepals,  the  outer 
stamens  are  converted  into  carpels,  these  supernumerary  and 
in  the  fruit  superposed  to  the  five  normal  carpels.2  In  Poppies, 
many  of  the  innermost  stamens  are  occasionally  transformed 
into  as  many  small  and  stalked  simple  pistils,  surrounding  the 
base  of  the  large  compound  one. 

1  It  must  not  be  concluded  that  the  supernumerary  petals  in  all  such  cases 
are  reverted  stamens,  or  stamens  and  pistils.  Some  are  instances  of  abnormal 
pleiotaxy,  i.  e.  of  the  production  of  one  or  more  additional  ranks  of  petals 
(better  deserving  the  name  of  double  flower},  with  or  without  reversion  of 
essential  organs  to  flower-leaves. 

8  These  trees  are  popularly  supposed  to  bear  fruit  without  blossoming; 
the  reverted  green  petals  being  so  inconspicuous  that  the  flower  is  un- 
noticed. 

FIG.  320,  321 .  Green  leaves  from  the  centre  of  a  blossom  of  Double-flowering  Cherry, 
one  still  showing,  by  its  partial  involution  and  its  style-like  apex,  that  it  is  a  reverted 
carpel,  the  other  a  small  but  well-formed  leaf. 

FIG.  322.    Leaf  or  leaflet  of  Bryophyllum,  developing  plantlets  along  the  margins. 


ITS  METAMOKPHY. 


173 


312.  Another  line  of  teratological  evidence  is  furnished  by 
prolification.     The   parts  of  the  flower  are,   by  the  doctrine, 
homologous  with  leaves,  and  no  leaf  ever  terminates  an  axis. 

Normally,  in  fact,  the 
axis  is  never  prolonged 
beyond  the  flower,  but 
abnormally  it  may  be. 
It  may  resume  vegeta- 
tive growth  as  a  termi- 
nal growing  bud,  either 
from  between  the  pistils 
after  the  whole  flower  is 
formed,  or  at  an  earlier 
period,  usurping  the 
central  part  of  the 
flower.  Thus,  when  a 
384  rose  is  borne  on  a  pe- 

duncle rising  from  the  centre  of  a  rose,  which  is 
not  very  unusual,  or  a  leafy  stem  from  the  top 
of  a  pear  (Fig.  323),  the  flower  was  probably 
complete  before  the  monstrous  growth  set  in.  In 
Fig.  324,  the  reversion  to  foliaceous  growth  took 
effect  after  the  stamens  but  before  the  pistils 
were  formed.  In  rose-fauds  out  of  roses,  the  terminal  proliferous 
shoot  takes  at  once  the  form  of  a  peduncle  ;  in  the  shoot  from 
the  pear,  that  of  a  leafy  stem. 

313.  Again,  axillary  buds  are  normally  formed  in  the  axil  of 
leaves.     No  such  branching  is  known  in  a  normal  flower.     But 
in  rare   monstrosities  a  bud   (mostly  a 

flower-bud)  makes  its  appearance  in  the 

axil  of  a  petal  or  of  a  stamen ;   and  it 

may  be   clearly  inferred  that  the  organ 

(not  itself  axillary)    from  the   axil  of 

which  a  bud  develops  is   a  leaf  or  its 

homologue.      Fig.  325  exhibits  a  clear 

case  of  the  kind,  a  flower  in  the  axil 

of  each  petal  of  Celastrus  scandens.     Flowers,  or  pedunculate 

clusters  of  flowers,  from  the  axil  of  petals  of  garden  Pinks  are 

sometimes  seen.     A  long-pedunculate  flower  from  the  axil  of  a 

FIG.  323.    A  monstrous  pear,  prolonged  into  a  leafy  branch  ;  from  Bonnet. 

FIG.  324.  Retrograde  metamorphosis  of  a  flower  of  the  Fraxinella  of  the  gardens, 
from  Lindley's  Theory  of  Horticulture ;  an  internode  elongated  just  above  the  stamens, 
and  bearing  a  whorl  of  green  leaves. 

FIG.  325.  A  flower  of  False  Bittersweet  (Celastrus  scandens)  producing  other 
flowers  in  the  axils  of  the  petals ;  from  Turpin. 


174  THE  FLOWER. 

stamen  of  a  species  of  Water-Lily  (Nymphaea  Lotus)  is  figured 
and  described  by  Dr.  Masters.1 

314.  In  the  application  of  morphological  ideas  to  the  elucida- 
tion of  the  flower,  nothing  should  be  assumed  in  regard  to  it 
which  has  not  its  proper  counterpart  and  exemplar  in  the  leaves 
and  axis  of  vegetation. 


SECTION  II.    FLORAL  SYMMETRY. 

315.  The  parts  of  a  flower  are  symmetrically  arranged  around 
its  axis.2    Even  when  this  symmetry  is  incomplete  or  imperfect,  it 
is  still  almost  always  discernible ;  and  the  particular  numerical 
plan  of  the  blossom  may  be  observed  or  ascertained  in  some  of 
the  organs. 

316.  Adopting  the  doctrine  that  the  parts  of  the  flower  are 
homologous  with  leaves,  the  sj^mmetry  is  a  consequence  of  the 
phyllotaxy.     It  is  symmetry  around  an  axis,  not  the  bilateral 
symmetry  which  prevails  in  the  animal  kingdom.     For  parts  of 
a  flower  disposed  in  a  continuous  spiral  (which  mostly  occurs 
when  they  are  numerous) ,  the  arrangement  is  that  of  some  order 
of  this  kind  of  phyllotaxy,  which  distributes  the  parts  equably 
into  superposed  ranks.     (237.)    The  much  commoner  case  of 


1  The  fullest  enumeration  and  discussion  of  the  very  various  kinds  of 
abnormal  structures  and  deviations  in  plants  is  to  be  found  in  the  Teratology 
of  Dr.  Masters,  above  referred  to.     Many  technical  terms  are  here  brought 
into  use,  which  need  not  be  here  mentioned,  except  the  following,  which  relate 
directly  to  floral  metamorphosis. 

Phyllody  (called  Phyllomorphy  by  Morren,  Frondescence  by  Engelmann)  is 
the  condition  wherein  true  leaves  are  substituted  for  some  other  organs; 
i.e.,  where  other  organs  are  metamorphosed  into  green  leaves.  There  is 
phyllody  of  pistils,  ovules,  filaments,  anther,  petals,  sepals,  &c. 

Sepcdody,  where  other  organs  assume  the  appearance  of  green  sepals. 

Petalody,  where  they  assume  the  appearance  of  petals,  as  normally  in 
Pinckneya  and  Calycophyllum,  in  which  one  calyx-lobe  enlarges  and  becomes 
petal-like,  and  abnormally  in  Primroses  where  all  the  calyx-lobes  imitate 
lobes  of  the  corolla  (this  has  been  termed  Calycanthemy) ;  also  of  the  stamens 
of  common  "  double  flowers." 

Staminody,  where  other  organs  develop  into  stamens.  Cases  of  this  as 
affecting  pistils  are  referred  to  above :  rarely  sepals  and  petals  are  so  affected. 

Pistillody,  where  other  organs  develop  into  pistils,  which  most  rarely 
happens  except  with  the  stamens,  as  above  mentioned. 

2  It  is  stated  that  Correa  de  Serra  (who  published  botanical  and  other 
papers  in  London,  Paris,  and  Philadelphia  during  the  first  twenty  years  of 
the  century,  but  who  knew  far  more  than  he  published)  was  the  first  botanist 
to  insist  on  the  symmetry  of  the  flower.     It  was  first  made  prominent  by  De 
Candolle,  in  the  The'orie  Ele'mentaire,  and  elaborated  in  detail  by  A.  St. 
Hilaire  in  his  Morphologic  Ve'ge'tale. 


FLOKAL  SYMMETRY.  175 

equal  number  of  parts  in  a  cycle,  and  the  cycles  alternating 
with  each  other,  is  simply  that  of  verticillate  phyllotaxy.  (234.) 
In  either  case,  the  members  of  the  successive  circles  (or  of 
closed  spirals  as  the  case  may  be)  will  be  equal  in  number ;  that 
is,  the  flower  will  be  isomerous. 

317.  A  Symmetrical  Flower  is  one  in  which  the  members  of  all 
the  cycles  (whorls  or  seeming  whorls)  are  of  the  same  number.1 
In  nature,  the  symmetry  is  of  all  degrees  :  it  is  most  commonly 
complete  and  perfect  as  to  the  floral  envelopes  when  it  is  not 
so  as  respects  the  essential  organs.     The  general  rule  is  that 
the  successive  cycles  alternate,  as  is  the  nature  of  true  whorls. 
But  the  superposition  of  successive  parts  is  not  incompatible 
with  symmetry  of  the  blossom,  although  it  is  a  departure  from 
the  ordinary  condition,  assumed  by  botanists  as  the  type.     An 
isomerous  flower  (meaning  one  with  an  equal  number  of  mem- 
bers of  all  organs)  is  the  same  as  symmetrical,  if  the  reference  be 
to  the  number  in  the  circles,  rather  than  to  the  total  number  of 
organs  of  each  kind.  % 

318.  A  Regular  Flower  is  one  which  is  symmetrical  in  respect 
to  the  form  of  the  members  of  each  circle,  whatever  be  their 
number  ;  i.  e. ,  with  the  members  of  each  circle  all  alike  in  shape. 

319.  These  two  kinds  of  symmetry  or  regularity,  with  their 
opposites  or  departures  from  symmetry,  need  to  be  practically 
distinguished  in  succinct  language.     For  the  terminology,  it  is 
best  to  retain  the  earlier  use,  generally  well  established  in  phyto- 
graphy,  as  above  defined. 

320.  A  Complete  Flower  is  one  which  comprises  all  four  or- 
gans, viz.  calyx,  corolla,  stamens,  pistil. 


1  This  is  not  only  the  definition  "  generally  applied  in  English  text-books," 
but  that  introduced  by  DeCandolle,  adopted  by  St.  Hilaire,  and  followed  at 
least  by  the  French  botanists  generally.  The  innovating  German  definition, 
of  a  recent  date,  is  that  a  symmetrical  flower  is  one  "  that  can  be  vertically 
divided  into  two  halves  each  of  which  is  an  exact  reflex  image  of  the  other." 
But  such  have  immediately  to  be  distinguished  into  "  flowers  which  can  be 
divided  in  this  manner  by  only  one  plane,"  which  Sachs  terms  "  simply 
symmetrical  or  monosymmetrical,"  and  those  which  can  be  symmetrically 
divided  by  two  or  more  planes,  "  doubly  symmetrical  or  poll/symmetrical," 
as  the  case  may  be.  Now  both  these  forms  have  a  more  expressive  and 
older  terminology,  adopted  by  Eichler,  viz. :  — 

Zygomorphous,  for  flowers,  or  other  structures,  which  can  be  bisected  in 
one  plane,  and  only  one,  into  similar  halves  (median  zygomorphous,  when 
this  is  a  median  or  antero-posterior  plane,  as  it  most  commonly  is ;  tram- 
verse  zygomorphous,  when  the  plane  of  section  is  transverse  or  at  right 
angles  to  the  median,  as  in  Dicentra) ; 

Actinomorphous  for  flowers,  &c.,  which  can  be  bisected  in  two  or  more 
planes  into  similar  halves. 


176  THE   FLOWEB. 

321.  Numerical  ground-plan.     Many  flowers  are  numerically 
indefinite  in  some  or  most  of  their  kinds  of  members,  as  Ranun- 
culus, Magnolia,  and  the  Rose  for  stamens  and  pistils,  Nym- 
phaea  for  all  but  perhaps  the  sepals,  many  Cactacese  for  all  but 
the  pistil,  and  Calycanthus  for  all  four  components.     But  more 
commonly  each  flower  is  constructed  upon  a  definite  numerical 
ground-plan  ;  and  the  number  is  usually  low.     Seldom,  if  ever, 
is  it  reduced  to  unity  in  a  hermaphrodite  blossom  (even  Hip- 
puris,  with  a  single  stamen  and  a  single  pistil,  is  not  an  un- 
equivocal case) ,  and  probably  never  in  a  complete  one.    But  there 
are  such  extremely  simplified  flowers  among  those  of  a  single  sex. 
In  Monocotyledons,  the  almost  universal  number  is  three,  some- 
times two  ;  in  ordinaiy  Dicotyledons,  five  prevails  ;  four  and  two 
are  not  uncommon  ;  three  is  occasional ;  and  higher  numbers  are 
not  wanting,  as  twelve  or  more  in  Houseleeks. 

322.  To  designate  the  particular  plan,  such  familiar  terms  of 
Latin  derivation  as  binary,  ternary,  quaternary,  quinary,  senary, 
&c.,  are  sometimes   employed,  denoting  that  the  parts  of  the 
flower  are  in  twos,  threes,  fours,  fives,  or  sixes.    More  technical 
and  precise  terms,  equivalent  to  these,  are  composed  of  the  Greek 
numerals  prefixed  to  the  word  meaning  parts  or  members,  as 

Monomerous,  for  the  case  of  a  flower  of  one  member  of  each  ; 

Dimerous,  of  two,  or  on  the  plan  of  two  members  of  each  ; 

Trimerous,  of  three,  or  on  the  plan  of  three  members ; 

Tetramerous,  of  four,  or  on  the  quaternary  plan  ; 

Pentamerous,  of  five,  or  on  the  quinary  plan  ; 

Hexamerous ,l  of  six,  or  on  the  plan  of  six  members  to  each 
circle.  But,  in  Monocotyledons,  so-called  hexamerous  blossoms 
are  really  trimerous,  the  sixes  being  double  sets  of  three. 

323.  Pattern  Flowers.     These  should  be  symmetrical,  regular, 

complete  in  all  the  parts  and  without  ex- 
cess or  complication  of  these,  and  with- 
out any  of  the  cohesions  or  adhesions 
( which  may  obscure  the  type,  or  render  it 
'  less  expressive  of  the  idea  that  a  flower 
consists  of  a  series  of  circles  or  spirals 
of  modified  leaves  crowded  on  a  short 
axis.  Wherefore  the  illustration  Fig.  307, 
with  its  diagram  Fig.  308,  may  serve  as  a  pattern  pentamerous 
or  quinary  flower;  and  Fig.  326,  with  its  diagram,  Fig.  327, 

1  These  may  be  shortly  written  1-merous,  2-raerous,  3-merous,  and  so  on  up 
to  10-merous  (decamerous) ,  12-merous  (dodecamerous) ,  &c. 

Fig.  326.  Parts  of  a  symmetrical  trimerous  flower  (Tillsea  muscosa):  a.  o*lyx; 
6.  corolla;  c.  stamens;  d.  pistils.  327.  Diagram  of  the  same. 


FLOEAL  SYMMETRY. 


177 


as  a  pattern  trimerous  or  ternary  flower;  these  being  simply 
isomerous,  and  of  one  circle  of  each  kind.  And  the  whole 
relation  of  the  parts,  viewed  as 
modified  leaves  on  the  common 
axis,  may  be  exhibited  in  such  a 
diagram  of  a  pattern  isostemonous 
5-merous  flower  as  that  displayed 
in  Fig.  328. 

324.  Diplostemonons  Type.   The 
foregoing    patterns    are    selected 
upon    the    idea  of   the    greatest 
simplicity   consistent    with    com- 
pleteness.    But  extended  observation  leads  to 
jkjp  the  conclusion  that  the  typical  flower  in  nature 

\V  A  has  two  series  of  stamens,  as  it  has  two  series  in 

the  perianth ;  that  is, 
as  many  stamens  as 
petals  and  sepals  taken 
together.1  As  the 
petals  alternate  with 
the  sepals,  so  the  first 
series  of  stamens  al- 
ternates with  the  pet- 
als, the  second  series 
of  stamens  alternates  with  the  first,  and  the  pistils  or  carpels 
when  of  the  same  number  alternate  with  these.  Thus  the  outer 
series  of  stamens  and  the  carpels  normally  stand  before  (are 


1  This  riew  of  the  symmetry  of  the  flower  was  first  taken  by  Brown 
(Obs.  PL  Oudney,  in  Denham  and  Clapperton  Trav.  1826,  reprinted  in  Ray 
Soc.  ed.  of  Collected  Works,  i.  293).  It  is  true  that  Brown  declares  the 
same  of  the  pistils  ;  but  that  is  not  made  out.  The  evidence  of  this  doctrine 
is  to  be  gathered  from  a  large  and  varied  induction ;  from  the  general  pres- 
ence of  the  two  sets  of  stamens,  and  no  more,  in  petaloideous  Monocotyle- 
dons ;  the  unaltered  position  of  the  carpels  (before  the  sepals)  when  the 
inner  set  of  stamens  is  wanting,  as  in  the  Iris  Family ;  the  very  common 
appearance  in  haplostemonous  flowers  among  the  Dicotyledons  of  vestiges 
of  a  second  series,  or  of  bodies  which  may  be  so  interpreted.  The  androecium 
or  the  blossom  is  said  to  be 

Isostemonous  or  Haplostemonous  when  the  stamens  are  of  one  series,  equal 
in  number  to  that  of  the  ground-plan  of  the  blossom ; 

Diplostemonous,  when  there  are  two  series,  or  double  this  number. 

FIG.  328.  Ideal  plan  of  a  plant,  with  the  simple  stem  terminated  by  a  symmetrical 
pentamerous  flower ;  the  different  sets  of  organs  separated  to  some  distance  from  each 
other,  to  show  the  relative  situation  of  the  parts.  One  of  each,  namely,  a,  a  sepal,  b, 
a  petal,  c,  a  stamen,  and  d,  a  pistil,  also  shown,  enlarged. 

FIG.  329.    A  pentamerous  diploatemonou*  flower  of  Sedum. 


178  THE    PLOVER. 

superposed  to)  the  sepals,  and  the  stamens  of  the  inner  series 
stand  before  the  petals  ;  as  in  the  diagram,  Fig.  33 1.1 

325.  Flowers  which  completely 
exemplify  their  type  or  symmetry 
are  rare,  but  most  exhibit  it  more 
or  less.  Each  natural  order  or 
group  exhibits  its  own  particular 
floral  type,  or  modification  of  the 
common  type.2  Some  of  these 
modifications  do  not  at  all  affect 
the  symmetry  or  obscure  the  plan 
of  the  flower,  except  by  combina- 
tions which  render  the  phylline 
character  of  the  floral  envelopes  and  carpels 
apparent,  such  combinations  being  of 
rare  occurrence  in  foliage.  Others  gravely 
interfere  with  floral  symmetry,  sometimes  to 
such  degree  that  the  true  plan  of  the  blossom 
is  to  be  ascertained  only  through  extended 
comparisons  with  the  flowers  of  other  plants 
of  the  same  order  or  tribe,  or  of  related 
orders.  The  symmetry  of  the  blossom  finds  its  explanation  in 
the  laws  which  govern  the  arrangement  of  leaves  on  the  axis  ; 
that  is,  in  phyllotaxy.  The  deviations  from  symmetry  and  from 
typical  simplicity  have  to  be  explained,  and  in  the  first  instance 


1  For  convenient  reference  and  the  avoidance  of  circumlocution,  some 
writers  term  the  stamens  which  are  before  the  petals  epipetalous,  those  before 
the  sepals  episepalous ;  but,  as  this  prefix  means  upon,  it  is  better  to  restrict 
these  terms  to  cases  of  adnation  of  stamens  to  these  respective  parts  of  the 
perianth,  and  to  distinguish  as 

Antipetalous,  those  stamens  which  stand  before  petals,  whether  adnate  or 
free,  and 

Antisepalous,  those  which  stand  before  sepals.  —  These  terms  we  find  have 
already  been  employed  in  this  way  by  Dr.  A.  Dickson  (in  Seemann,  Jour. 
Bot.  iv.  275),  with  the  addition  of  a  third,  viz. 

Parapetalous,  for  stamens  which  stand  at  each  side  of  a  petal,  yet  not 
necessarily  before  a  sepal,  as  in  many  Rosaceae. 

2  These  particular  types,  with  their  modifications,  are  set  forth  in  the 
characters  or  distinguishing  marks  of  the  orders,  tribes,  genera,  &c.     The 
best  generally  available  illustrations  of  ordinal  types  are  in  Le  Maout  and 
Decaisne's  Traite'  Ge'ne'ral  de  Botanique,  and  in   Hooker's  English  edition 
and  revision,  entitled  A  General  System  of  Botany,  Descriptive  and  Analyti- 
cal, London,   1873.     The   best   morphological   presentation  is  in  Eichler'g 
Bluthendiagramme,  &c.  (Flower  Diagrams,  Constructed  and   Illustrated), 
Leipzig,  1875. 

FIG.  330.    Opened  flower  of  Trillium  erectum.    331.    Diagram  of  the  same. 


ITS   GENERAL  MODIFICATIONS.  179 

to  be  classified.  To  have  morphological  value,  such  explanation 
should  be  based  upon  just  analogies  in  the  foliage  and  other 
organs  of  vegetation.  Whatever  is  true  of  leaves  and  of  the 
vegetating  axis  as  to  position  of  parts,  mode  of  origin  and 
growth,  division,  connection,  and  the  like,  may  well  be  true  of 
homologous  organs  in  the  flower. 

SECTION  III.     VARIOUS  MODIFICATIONS  OF  THE  FLOWER. 
§  1.     ENUMERATION  OF  THE  KINDS. 

326.  In  the  morphological  study  of  flowers,  these  modifica- 
tions are  viewed  as  deviations  from  type.     Their  interpretation 
forms  no  small  part  of  the  botanist's  work.     They  may  be  classed 
under  the  following  heads  :  — 

1.  Union  of  members  of  the  same  circle  :  COALESCENCE. 

2.  Union  of  contiguous  parts  of  different  circles  :  ADNATION. 

3.  Inequality  in  size,  shape,  or  union  of  members  of  the  same 
circle :  IRREGULARITY. 

4.  Non-appearance  of  some  parts  which  are  supposed  in  the 
type  :  ABORTION  or  SUPPRESSION. 

5.  Non-alternation   of  the   members   of  contiguous   circles : 
ANTEPOSITION  or  SUPERPOSITION. 

6.  Increased   number  of  organs,   either  of  whole  circles  or 
parts  of  circles  :  AUGMENTATION  or  MULTIPLICATION. 

7.  Outgrowths,  mostly  from  the  anterior  or  sometimes  pos- 
terior face  of  organs  :  ENATION. 

8.  Unusual  development  of  the  torus  or  flower- axis. 

9.  To  which  may  be  appended  morphological  modifications, 
some  referable  to  these  heads  and  some  not  so,  which  are  in 
special  relation  to  the  act  of  fertilization.     These  are  specially 
considered  in  Section  IV. 

327.  These  deviations  from  assumed  pattern  are  seldom  single  ; 
possibly  all  may  coexist  in  the  same  blossom.     Several  of  them 
occur  even  in  that  one  of  the  orders,  the  Crassulacese,  which 
most  obviously  exhibits  the  normal  type  throughout. 

328.  Thus,  Sedum  (Fig.  329),  with  two  circles  of  stamens, 
being  taken  as  the  true  type  (324),  Crassula  (Fig.  307)  wants 
the  circle  of  stamens  before  the  petals  ;  Tillsea  (Fig.  326)  is  the 
same,  but  with  the  members  symmetrically  reduced  from  five  to 
three ;  Rhodiola  loses  all  the  stamens  by  abortion  in  one  half 
the  individuals  and  the  pistils  in  the  other,   sterile  rudiments 
testifying  to  the   abortion ;   Triactina  has  lost  two  of  its  five 
carpels,  and  the  three  remaining  coalesce  into  one  body  up  to  the 


180  THE  FLOWER. 

middle ;  Penthorum  (Fig.  335,  336)  has  its  five  carpels  coales- 
cent  almost  to  the  top,  and  usually  loses  its  petals  by  abortion  ; 
in  Grammanthes  and  Cotyledon  (Fig.  332-334),  the  sepals  are 
coalescent  into  a  cup  and  the  petals  into  a  deeper  one,  out  of 


which  the  stamens  appear  to  arise,  these  being  adnate  to  the 
corolla.  Symmetrical  increase  in  the  number  of  members  of 
each  circle  is  no  proper  deviation  from  type,  at  least  in  this 
family  (in  which  flowers  on  the  same  plant  sometimes  vary  from 
5-merous  to  4-merous  and  6-merous)  ;  and  in  Sempervivum  (to 
which  Houseleek  belongs)  these  members  are  always  more  than 
five  and  sometimes  as  many  as  twenty  in  each  circle. 


§  2.     REGULAR  UNION  OF  SIMILAR  PARTS. 

329.  Coalescence,  or  the  cohesion  by  the  contiguous  margins 
of  parts  of  the  same  circle  or  constituent  set  of  organs,  is  so  fre- 
quent that  few  flowers  are  completely  free  from  it.     The  last 
preceding  figures  show  it  in  the  gyncecium  and  corolla.     Fig. 
471-476  further  illustrate  it  in  the  corolla,  and  in  various  degrees 
up  to  entire  union  ;  and  Fig.  483-488  illustrate  it  in  the  andros- 
cium.     The   technical  terms  which  coalescence  calls  for,  and 
which  are  needful  in  botanical  description,  may  be  found  under 
the  account  of  the  particular  organ,  and  in  the  Glossary.     Such 
growing  together  of  contiguous  members  in  the  blossom  is  strictly 
paralleled  by  connate-perfoliate  leaves  of  ordinary  foliage  (212, 
Fig.  215),  where  it  more  commonly  occurs  in  upper  leaves,  and 
in  bracts,  which  are  still  nearer  the  flower. 

330.  It  should  now  be  hardly  necessary  to  explain  that  the 
terms  coalescence,  cohesion,  union,  and  the  corresponding  phrases 


FIG.  332.  Flower  of  Grammanthes.  333.  Flower  of  a  Cotyledon.  334.  The  corolla 
laid  open  showing  the  two  rows  of  stamens  inserted  on  It.  335.  The  five  pistils  of 
Penthorum,  united,  336.  A  cross-section  of  the  same. 


UNION  OF  PARTS.  181 

in  the  next  paragraph,  do  not  mean  that  the  parts  were  once  sepa- 
rate and  have  since  united.  That  is  true  only  of  certain  cases.  The 
union  is  mostly  congenital,  equally  so  in  the  disks  of  foliage  of 
the  Honeysuckle  (Fig.  215)  and  in  the  corolla  of  a  Convolvulus. 
The  lobes  which  answer  to  the  tips  of  the  constituent  leaves  of 
the  cup  or  tube  are  usually  first  to  appear  in  the  forming  bud, 
the  undivided  basal  portion  comes  to  view  later.  It  might  be 
more  correct  to  say  that  the  several  leaves  concerned  have  not 
isolated  themselves  as  they  grew.  Accordingly,  Dr.  Masters 
would  substitute  for  coalescence  and  adnate  the  term  inseparate. 
But  the  common  language  of  morphology  needs  no  change,  as  it 
consistently  proceeds  on  the  idea,  and  the  prevalent  fact,  that 
leaves  are  separate  things,  and  that  the  tube,  cup,  or  "  insepa- 
rate "  base  of  a  calyx  or  corolla,  consists  of  a  certain  number  of 
these.  It  is  no  contradiction  to  this  view  that  they  developed 
in  union.1 

§  3.    UNION  OF  DISSIMILAR  OR  SUCCESSIVE  PARTS. 

331.  Adnation  is  the  most  appropriate  term  to  denote  the  organic 
and  congenital  cohesion  or  consolidation  of  different  circles,  the 

1  If  it  were  seriously  proposed  to  change  the  language  of  descriptive 
botany  in  this  regard,  consistency  would  require  its  total  reconstruction,  with 
the  abolition  of  all  such  terms  as  cleft,  parted,  &c. ;  for  the  structures  in 
question  are  no  more  cleft  than  they  are  united.  While  these  convenient 
and  long-familiar  terms  are  continued  in  use  (as  they  surely  will  be),  although 
quite  contrary  to  literal  fact,  it  cannot  be  amiss  to  continue  those,  such  as 
connate,  adnate,  coalescent,  &c.,  which  imply  and  suggest  the  fundamental  fact 
in  the  structure  of  phaenogamous  and  the  higher  cryptogamous  plants,  viz. 
that  leaves  are  normally  unconnected  organs. 

Whether  fusion  or  separation  is  the  more  complex  condition,  and  therefore 
indicative  of  higher  rank,  is  a  question  of  a  different  order.  It  is  argued 
that  the  fusion  or  lack  of  separation  is  an  arrest  of  development,  and  there- 
fore an  indication  of  low  rank  or  less  perfection  than  the  contrary.  But  a 
phylogenetic  view  of  the  whole  case  may  reverse  this  conclusion  as  respects 
the  blossom.  The  course  of  development  from  thallus  and  frond  to  distinct 
foliage  on  an  axis,  from  little  to  full  differentiation,  is  clearly  a  rise  in 
rank,  as  also  is  the  differentiation  of  foliage  into  ordinary  leaves,  petals, 
stamens,  and  pistils.  But  there  is  as  much  differentiation  in  the  flower 
of  a  Convolvulus  as  of  a  Ranunculus,  and  more  in  that  of  a  Salvia,  a 
Lobelia,  and  an  Orchis.  In  all  such  flowers,  the  combination,  the  irregu- 
larity, and  the  diversification  in  many  cases  of  the  members  of  the  same  circle, 
all  indicate  complexity,  greater  specialization,  and  therefore  higher  rank. 
The  production  of  leaves  distinct  from  the  axis  is  one  step  in  the  ascending 
scale  :  such  specializations  and  combinations  of  these  as  occur  in  flowers  are 
higher  steps ;  and  the  most  specialized,  complex,  and  therefore  highest  in 
rank  are  complete,  corolliferous,  irregular  flowers,  with  a  definite  number  of 
members,  and  these  combined  in  view  of  the  adaptations  by  which  the  ends 
of  fertilization  and  fructification  are  best  subserved. 


182 


THE   FLOWER. 


apparent  growing  of  one  part  on  or  out  of  another,  —  as  of 
the  corolla  out  of  the  calyx,  the  stamens  out  of  the  corolla,  or 
all  of  them  out  of  the  pistil.  This  disguises  the  real  origin 
of  the  floral  organs  from  the  receptacle  or  axis,  in  successive 
series,  one  within  or  above  the  other.  Organs  in  this  condition 
are  also  and  rightly  said  to  be  Connate  (born  united)  ;  but,  as 
this  term  is  equally  applicable  to  the  coalescence  of  members  of 
the  same  circle,  the  word  Adnate  is  preferable,  as  applying  to 
the  present  case  only.  Adnation  is  heterogeneous  organic  co- 
hesion or  adhesion :  coa- 
lescence is  homogeneous 
cohesion  or  union. 

332.  Adnation  occurs  in 
very  various  degrees,  and 
affects  either  some  or  all  the 
organs  of  the  flower.  Its 
consideration  introduces  into 
terminology  several  peculiar 
terms,  which  may  here  be 
defined  in  advance.  Three 
of  them,  introduced  and 
prominently  employed  by 
Jussieu,  depend  upon  the 
degree  of  adnation,  or  the 
absence  of  it,  viz. :  — 

Hypogynous  (literally  be- 
neath pistil) ,  applied  to  parts 
which  are  inserted  (i.  e.  are 
borne)  on  the  receptacle  of 
the  flower,  as  in  Fig.  336. 
This  is  the  absence  of 
adnation,  or  the  condition 
which  corresponds  with  the 
unmodified  type. 

Perigynous  (around  the 
pistil)  implies  an  adnation 
which  carries  up  the  inser- 
tion of  parts  (which  always  means  apparent  origin  or  place  of 
attachment)  to  some  distance  above  or  away  from  the  recep- 

FIG.  336°.  Vejtical  section  of  a  flower  of  the  Common  Flax,  showing  the  normal  or 
hypogynous  insertion  of  parts  upon  the  torus  or  receptacle. 

FIG.  337.  Vertical  section  of  a  flower  of  the  Cherry,  to  show  the  perigynous  insertion, 
or  adnation  to  the  calyx,  of  the  petals  anil  stamens. 

FIG.  338.  Similar  section  of  the  flower  of  the  Purslane,  showing  an  adnation  of  all 
the  parts  with  the  lower  half  of  the  ovary. 


UNION    OF   DISSIMILAR    PAETS. 


183 


tacle,  so  commonly  placing  this  insertion  around  instead  of 
beneath  the  pistil;  whence  the  name.  The  perigyny  may  be, 
as  the  figures  show,  merely 
the  adnation  of  petals  and 
stamens  to  calyx,  the  calyx 
remaining  hypogynous,  as  in 
Fig.  337;  or  else  the  adna- 
tion of  the  calyx,  involving 
the  other  organs,  to  the  lower 
part  of  the  ovary,  as  in  Fig. 
338,  or  up  to  the  summit  of 
the  ovary,  while  the  petals 
and  stamens  are  adnate  still  further  to  the  calyx,  as  in  Fig.  339. 
The  latter  passes  into  what  is  called 

Epigynous  (on  the  pistil),  where  the  adnation  is  complete  to 
the  very  top  of  the  ovary,  and  none  beyond  it,  as  in  Fig.  340, 
341.  Yet  here  the  parts  so  termed  are  not  really  on  the  ovary, 
except  where  an  epigynous  disk  (394)  actually  surmounts  it. 


340  341 

333.  Adnation  brings  some  other  terms  into  use  in  botanical 
descriptions,  especially  those  of  superior  and  inferior.  In  this 
connection,  these  words  (in  Latin  taking  the  form  of  superus  and 
inferus)  denote  the  position  in  respect  to  each  other  of  ovary  and 
floral  envelopes,  —  not  the  morphological,  but  the  apparent  posi- 
tion or  place  of  origin.  Thus,  in  Fig.  336  and  in  337,  the  calyx 
is  inferior,  or  in  other  words  the  ovary  superior.  Here  real  and 
apparent  origin  agree,  this  being  the  normal  condition,  which 
is  otherwise  expressed  by  saying  that  the  parts  are/ree,  i.  e.  free 
from  all  adnation  of  one  to  the  other.  But,  in  Fig.  339-341,  the 

FIG.  339.  Similar  section  of  a  flower  of  Hawthorn,  showing  complete  adnation  to 
the  summit  of  the  ovary  and  of  the  other  parts  beyond. 

FIG.  340.  Vertical  section  of  a  Cranberry-flower,  and  341,  of  flower  of  Aralia 
nudicaulis.  with  so-called  epigynous  insertion  of  calyx,  corolla,  and  stamens;  the  calyx 
of  the  latter  completely  consolidated  with  the  surface  of  the  ovary,  or  its  limb 
obsolete. 


184  THE   FLOWER. 

ovary  is  said  to  be  inferior  and  the  calyx  superior,  the  calyx 
and  other  parts,  in  consequence  of  the  adnation  of  its  lower  part, 
seeming  to  rise  from  the  summit  of  the  ovary. 

334.  Adnation   of  floral  envelopes   to   pistil  rarely  extends 
beyond  the  ovary ;  yet,  in  species  of  Iris  having  a  tube  to 
the  perianth,  this  tube  is  commonly  adnate  for  most  of  its  length 
to  the  style.     But  when  the  calyx  has  its  tube  or  portion  with 
united  sepals  prolonged,  the  petals  and  the  stamens  are  usually 
adnate  more  or  less  to  it,  i.  e.  are  inserted  on  the  calyx.     And, 
when  the  petals  are  united  and  prolonged  into  a  tube,  the  sta- 
mens,   being  within   the   corolla,  are   commonly  adnate   to  or 
inserted  upon  this. 

335.  No  one  doubts  that  the  view  is  a  true  one  which  repre- 
sents the  perianth-tube  as  adnate  to  the  style  in  Iris,  petals  and 
stamens  as  adnate  to  calyx  in  the  Cherry  (Fig.  337),  stamens 
as  adnate  to  base  of  corolla  in  Fig.  334,  and  a  long  way  farther 
in  Phlox,  &G.     That  the  calyx  is  similarly  adnate  to  the  ovary 
is  nearly  demonstrable  in  certain  cases. 

336.  But,  as  the  lower  portion  of  a  pear  is  undoubtedly  recep- 
tacle, or  rather  the  enlarged  extremity  of  the  flower-stalk,  as  in  a 
rose  at  least  a  portion  of  the  hip  is  receptacle,  as  the  tube  of  the 
flower  in  a  Cereus  or  other  Cactacea  has  all  the  external  char- 
acters and  development  of  a  branch,  so  it  is  most  probable  that 
in  many  cases  the  supposed  calyx-tube  adnate  to  an  inferior 
ovary  is  partly  or  wholly  a  hollowed  receptacle  (in  the  manner 
of  a  Fig-fruit)  ;  that  is,  a  cup-shaped  or  goblet-shaped  develop- 
ment of  the  base  of  the  floral  axis.     This  would  bring  the  case 
under  §  7.  (326,  495.) 


§  4.    IRREGULARITY  OF  SIMILAR  PARTS. 

337.  Irregularity,  or  inequality  in  form  or  in  union  of  mem- 
bers of  a  circle,  is  extremely  common,  either  with  or  without 
numerical  symmetry.     One  or  two  examples  may  suffice. 

338.  Irregular  flowers  with  symmetrical  perfection,  except  in 
the  gynoscium,  are  well  seen  in  the  Pea  Family,  to  which  belongs 
the  kind  of  corolla  called  Papilionaceous,  from  some  imagined 
resemblance   to   a  butterfly.    (Fig.  342-344.)      This   flower  is 
5-merous  throughout,  has  the  full  complement  of  stamens  (10,  or 
two  sets) ,  but  the  gynoacium  reduced  to  a  single  simple  pistil. 
The  striking  irregularity  is  in  the  corolla,  the  petals  of  which 
bear  distinguishing  names  :  the  posterior  and  larger  one,  exter- 
nal in  the  bud,  is  the  VEXILLUM  or  STANDARD  (Fig.  344,  a) ; 


INEQUALITY   OF   SIMILAR   PARTS. 


185 


the  two  lateral  next  and  under  the  standard,  AJLM  or  WINGS  (b)  ; 
the  two  anterior,  covered  by  the  wings  and  partly  cohering  to 


form  a  prow-shaped  body  (c) ,  the  CARTNA  or  KEEL.  The  calyx 
is  slightly  irregular  by  unequal  union,  the  two  upper  sepals 
united  higher  than  the 
other  three.  The  sta- 
mens are  much  more 
coalescent,butwith  an 
irregularity,  nine  com- 
bined by  the  lower  part 
of  their  filaments,  and 
one  (the  posterior) 
separate.  (Fig.  345.) 
339.  The  plan  and 

floral  symmetry  in  the  Locust-blossom  and  347 
its  relatives  are  little  obscured  by  the  irregu- 
larities and  the  coalescence,  hardly  more  so 
than  in  the  plainer  flower  of  its  relative, 
Baptisia  (Fig.  347,  348),  in  which  the  petals  are  somewhat 
alike,  and  the  ten  stamens  are  distinct  or  unconnected.  Only 
the  calyx  is  more  irregular,  by  the  union  of  the  two  posterior 
sepals  almost  to  the  tip.  (Fig.  348.) 

FIG.  342.  Diagram  of  flower  of  the  Locust,  Robinia  Pseudacacia :  a.  axis  of  inflores- 
cence; b.  bract;  first  circle  of  5,  calyx;  five  remaining  pieces,  corolla;  next  anthers, 
10  in  number;  in  the  centre  a  single  simple  pistil.  343  Front  yiew  of  Locust-flower 
showing  only  the  corolla.  344.  This  corolla  displayed. 

FIG.  345.  Androecium  of  the  Locust,  nine  stamens  coalescent,  one  distinct.  346. 
Same  of  a  Lupine,  all  ten  filaments  coalescent  below  into  a  closed  tube. 

FIG.  347.  Calyx  and  corolla  of  Baptisia  australis.  348.  Same  with  petals  fallen, 
showing  ten  distinct  stamens  and  tip  of  the  style. 


186 


THE  FLO  WEE. 


<ie 


340.  But  in  a  Lupine-blossom,  of  equally  near  relationship, 
a  casual  observer  might  fail  to  recognize  the  very  same  type, 
although  disguised  only  by  cohesions.  For  while  the  two  pos- 
terior sepals  are  united  to  the  tip  on  one  side  of  the  blossom,  the 
three  others  are  similarly  united  into 
one  body  on  the  anterior  side,  giving 
the  appearance  of  two  sepals  instead 
of  five :  in  the  corolla,  the  two  keel- 
petals  are  more  strictly  united  into  a 
slender  scythe-shaped  or  sickle-shaped 
body ;  so  that  the  petals  might  with 
the  unwary  pass  for  four:  in  the 
androecium,  the  coalescence  includes  all 
ten  stamens  (Fig.  346),  which  is  an 
approach  to  regularity. 

341.  The  5-merous  symmetry  of  the 
Violet-blossom  is   complete  until  the 
gyncecium  is  reached  (but  with  only 
one  circle  of  stamens)  ;  the  main  irregu- 
larity of  the  perianth  is  in  the  anterior 
petal,  with  its  nectariferious  sac  at  base 
(Fig.  349-351)  ;  the  two  stamens  near- 
est this  send  into  the  sac  curious  appendages,  which  the  other 
three  do  not  possess  ;  the  gynoecium  is  composed  of  three  car- 
@  pels  coalescent  into  one  compound  ovary  in  a 

^^fflii  manner  hereafter  explained.     In  Antirrhinum 

>^  and  Linaria  (Fig.  480,  481),  there  is  a  similar 
irregularity  accompanying  coalescence  of  the 
^  petals,  the  anterior  one  being  extended  at  base 
into  a  nectariferous  sac  or  hollow  spur.1  The 
flower  of  a  Lobelia  (Fig.  488)  has  the  same 
numerical  plan  and  symmetry  as  that  of  Viola 
(except  that  the  gyno3cium  is  dimerous)  ;  but 
members  are  adnate  below  and  coalescent  above,  and  the 
corolla  is  irregular  through  unequal  coalescence  of  the  five  petals, 
and  the  absence  of  coalescence  down  one  side. 

1  PELORIA  is  a  name  given  by  Linnaeus  to  an  occasional  monostrosity  of 
these  flowers  (imitated  in  sundry  others),  in  which  the  base  of  every  petal,  or 
answering  part  of  the  corolla,  is  prolonged  downward  into  a  sac  or  spur. 
The  sac  is,  morphologically  considered,  a  departure  from  normal  regularity  : 
in  the  monster,  symmetrical  regularity  is  restored  by  the  development  of  four 
more  sacs. 

FIG.  349.  Flower  of  Viola  sagittata.  350.  Its  sepals  and  petals  displayed.  351. 
Diagram  of  a  Violet-blossom,  from  Eichler,  with  bract  or  subtending  leaf  (below),  a  pair 
of  bractlets  (lateral),  and  axis  to  which  the  subtending  leaf  belongs  (above  or  posterior). 


DISAPPEARANCE  OF   PARTS.  187 


§  5.  DISAPPEARANCE  OR  OBLITERATION  OF  PARTS. 

342.  Abortion  or  Suppression  are  somewhat  synonymous  terms 
to  denote  the  obliteration  or  rather  non-appearance  of  organs 
which  belong  to  the  plan  of  the  blossom.     Abortion  is  applied 
particularly  and  more  properly  to  partial  obliteration,  as  where 
a  stamen  is  reduced  to  a  naked  filament,  or  to  a  mere  rudiment 
or  vestige,  answering  to  a  stamen  and  occupying  the  place  of 
one,  but  incapable  of  performing  its  office  ;  suppression,  to  abso- 
lute non-appearance.     Such  vestiges  or  abortive  organs  justify 
the  use  of  these  terms,  the  more  so  as  all  gradations  are  some- 
times met  with  between  the  perfect  organ  and  the  functionless 
rudiment  which  occupies  its  place.     Such  obliterations,  whether 
partial  or  complete,  may  affect  either  a  whole  circle  of  organs 
or  merely  some  of  its  members.     The  former  interferes  with  the 
completeness  of  a  flower,  and  may  obscure  the  normal  order  of 
its  parts.     The  latter  directly  interferes  with  the  symmetry  of 
the  blossom,  and  is  commonly  associated  with  irregularity. 

343.  Of  parts  of  a  Circle.      Among  papilionaceous   flowers 
(338) ,  different  species  of  Erythrina  have  all  the  petals  but  one 
(the  vexillum,  Fig.  344,  a)  much  reduced  in 

size,  in  some  concealed  in  the  calyx,  and  in 
every  way  to  be  ranked  as  abortive  organs. 
In  Amorpha,  of  the  same  family,  these  four 
petals  are  gone,  leaving  no  trace,  reducing 
the  corolla  to  a  single  petal.  (Fig.  352,  353.) 
This  one  is  evidently  the  vexillum,  both  by 
position  and  shape  ;  and  the  5-merous  type, 
also  the  particular  type  of  the  family,  are  still 
discernible  in  the  five  notches  of  the  calyx, 
the  ten  stamens,  &c.  In  a  related  genus, 
Parryella,  even  this  last  petal  is  wanting,  and 
the  andrcecium  is  straight,  all  irregularity  thus 
disappearing  through  suppression. 

344.  Delphinium   or  Larkspur  and   Aconite  or  Monkshood 
furnish  good  examples  of  flowers  in  which  irregularity  is  accom- 
panied by  more  or  less  abortion.     The  calyx  of  the  Larkspur 
(Fig.  354-356)  is  irregular  by  reason  of  the  dissimilarity  of  the 
five  sepals,  one  of  which,  the  uppermost  and  largest,  is  pro- 
longed posteriorly  into  a  long  and  hollow  spur.     Within  these, 
and  alternate  with  them  as  far  as  they  go,  are  the  petals,  only 

FIG.  352.    Stamens  and  pistil  of  Amorpha  fruticosa.    353.    An  entire  flower  of  the 
same. 


188 


THE  FLOWER. 


four  in  number,  and  these  of  two  shapes,  the  two  upper  ones 
having  long  spurs  which  are  received  into  the  spur  of  the  upper 
the  two  lateral  ones  having  a  small  but  broad  blade 


raised  on  a  stalk-like  claw ;  and  the  place  which  the  fifth  and 
lower  petal  should  occupy  (marked  in  the  ground-plan,  Fig.  356, 


PIG.  354.  Flower  of  a  Larkspur.  355.  The  five  sepals  (outer  circle)  and  the  four 
petals  (inner  circle)  displayed.  356.  Ground-plan  of  the  calyx  and  corolla. 

FIG.  357.  Flower  of  an  Aconite  or  Monkshood.  358.  The  five  sepals  and  the  two 
small  and  curiously  shaped  petals  displayed ;  also  the  stamens  and  pistils  in  the  centre. 
359.  Ground-plan  of  the  calyx  and  corolla;  the  dotted  lines,  as  in  Fig.  356,  representing 
the  suppressed  parts. 


DISAPPEARANCE   OF   PARTS.  189 

by  a  short  dotted  line)  is  vacant,  this  petal  being  suppressed, 
thereby  rendering  the  blossom  unsymmetrical.  In  Aconite 
(Fig.  357-359),  the  plan  of  the  blossom  is  the  same, 
but  the  uppermost  and  largest  of  the  five  dissimilar 
sepals  forms  a  helmet-shaped  or  hood-like  body; 
three  of  the  petals  are  wanting  altogether  (their 
places  are  shown  by  the  dotted  lines  in  the  ground- 
plan,  Fig.  359)  ;  and  the  two  upper  ones,  which  ex- 
tend under  the  hood,  are  so  reduced  in  size  and  so 
anomalous  in  shape  that  they  would  not  be  recog- 
nized as  petals.  One  of  these,  enlarged,  is  exhibited 
in  Fig.  360.  Petals  and  other  parts  of  this  and  of  va- 
rious extraordinary  forms  were  termed  by  Linnaeus 
NECTARIES,  a  somewhat  misleading  name,  as  they 
are  no  more  devoted  to  the  secretion  of  nectar  than 
ordinary  petals  or  other  parts  are.  In  these  flowers, 
moreover,  the  stamens  are  much  increased  in  number.  m 

345.  Analogous  abortion  of  some  of  the  stamens,  along  with 
a  particular  irregularity  of  the  perianth,  especially  of  the  corolla, 
characterizes  a  series  of  natural  orders  with  coalescent  petals.1 
These  flowers  are  all  on  the  5-merous  plan  (except  that  the 
gyncecium  is  2-merous),  but  with  corolla,  and  not  rarely  the 
calyx,  irregular  through  unequal  union  in  what  is  called  the  bila- 
biate or  two-lipped  manner.  The  greater  union  is  always  median, 
or  anterior  and  posterior,  and  two  of  the  coalescent  members  form 
one  lip,  three  the  other.  The  two  posterior  petals  form  the 
upper  lip,  the  anterior  and  two  lateral  form  the  lower  lip  of  the 
corolla ;  in  the  calyx,  when  that  is  bilabiate,  this  is  of  course 
reversed.  In  some,  as  in  Sage  and  Snapdragon,  the  bilabiation 
of  the  corolla  is  striking  (Fig.  479-481),  and  readily  comparable 
to  the  two  jaws  of  an  animal ;  in  others,  the  parts  are  almost  regu- 
lar. The  suppression  referred  to  is,  in  most  of  these  cases,  that 
of  the  posterior  of  the  five  stamens,  as  in  Fig.  361,  where  it  is 
complete.  In  Pentstemon  (Fig.  362),  a  sterile  filament  regu- 
larly occupies  the  place  of  the  missing  stamen.  The  position 
sufficiently  indicates  its  nature.  This  is  also  revealed  by  the 
rare  occurrence  of  an  imperfect  or  of  a  perfect  anther  on  this 


1  These  natural  orders  in  which  this  occurs,  or  tends  to  occur,  are  the 
Scrophulariaceae  (Snapdragon,  Pentstemon,  Mimulus,  &c.),  Orobanchaceae 
(Beech-drops),  Lentihulaceae  (Bladderwort),  Gesneraceae  (Gloxinia),  Big- 
noniaceae  (Trumpet  Creeper,  Catalpa),  Pedaliaceae  (Martynia),  Acanthaceae, 
Labiatae  (Salvia,  Stachys),  &c. 

FIG.  360.    A  petal  (nectary)  of  an  Aconite,  much  enlarged. 


190 


THE  FLOWEB. 


filament,  —  a  monstrosity,  indeed,  but  the  monstrosity  is  here 
a  return  to  normal  sj-mmetry.  The  two  stamens  nearest  the 
suppressed  or  abortive  one  generally 
share  in  the  tendency  to  abortion,  as 
is  shown  by  their  lesser  length  or 
smaller  anthers :  in  the  flower  of 
Catalpa,  these  two  also  are  either  im- 
perfect or  reduced  to  mere  vestiges 
(as  in  Fig.  363)  :  in  very  many  other 
plants  of  these  families,  even  these 
vestiges  are  not  seen,  and  so  the  five 
stamens  are  by  abortion  or  complete 
suppression  reduced  to  two. 

346.  Suppression  in  the  gynoecium 
to  a  number  less  than  the  numerical 
plan  of  the  flower  (as  shown  in  the 
perianth)  is  of  more  common  occur- 
rence than  the  typical  number,  and 
the  reduction  is  comparatively  con- 
stant throughout  the  genus  or  order. 
A  papilionaceous  or  other  leguminous 
flower  with  more  than  one  or  with  all 
five  pistils  is  exceedingly  rare,  and 
except  in  one  pentacarpellary  genus 
is  a  monstrosity.  Suppression  of 
the  interior  is  more  common  than 
of  exterior  organs.  Want  of  room 
in  the  bud  may  partially  explain 
this. 

347.  Suppression  of  whole  Circles.  Such  suppression  or  rather 
non-production  in  the  actual  blossom  of  whole  series  of  organs 
which  belong  to  the  type,  and  indeed  are  sometimes  present  in 
that  blossom's  nearest  relatives,  is  very  common.  It  gives 
occasion  to  several  descriptive  terms,  which  may  be  here  defined 
together.  First,  and  in  general,  flowers  are 

Incomplete,  in  which  any  one  or  more  of  the  four  kinds  of  organs 
is  wanting,  whatever  these  may  be  ; 

Apetalous,  when  the  corolla  or  inner  perianth  is  wanting ; 
Monochlamydeous,   where  the   perianth  is  simple  instead  of 

FIG.  361.  Corolla  of  Gerardia  purpurea  laid  open,  with  the  four  stamens,  the  place 
which  the  fifth  should  occupy  indicated  by  a  cross. 

FIG.  362.  Corolla  of  Pentstemon  grandiflorus  laid  open,  with  its  four  stamens,  and 
a  sterile  filament  in  the  place  of  the  fifth  stamen. 

FIG.  363.  Corolla  of  Catalpa  laid  open,  with  two  perfect  stamens  and  the  vestiges 
of  three  abortive  ones. 


DISAPPEARANCE  OF   PARTS.  191 

double,  in  which  case  the  wanting  set  is  generally  (but  not  quite 
always)  the  inner,  or  the  corolla  ; 

Dichlamydeous,  when  both  circles  of  the  perianth  (calyx  and 
corolla)  are  present ; 

Achlamydeous,  when  both  are  wanting,  as  in  Fig.  365.  (These 
three  terms  are  seldom  employed.) 

Unisexual  (also  Diclinous  or  Separated) ,  when  the  suppression 
is  either  of  the  stamens  or  the  pistils.  In  contradistinction,  a 
flower  which  possesses  both  is  Bisexual  or  Hermaphrodite. 

Staminate,  or  Male,  when  the  stamens  are  present  and  the 
pistils  absent ; 

Pistillate,  or  female,  when  the  pistils  are  present  and  the 
stamens  absent ; 

Monoecious  (of  one  household),  when  stamens  and  pistils  oc- 
cupy different  flowers  on  the  same  plant ; 

Dioecious  (of  two  households),  when  they  occupy  different 
flowers  on  different  plants  ; 

Polygamous,  when  the  same  species  bears  both  unisexual  and 
bisexual  or  hermaphrodite  flowers.  This  may  occur  in  various 
ways,  from  the  greater  or  less  abortion  of  either  sex,  either  on 
the  same  or  on  separate  individual  plants  ;  —  as  Monceciously  or 
Diceciously  Polygamous,  according  to  the  tendency  to  become  either 
monoecious  or  dioecious.  Recently  Darwin  has  well  distinguished 
the  case  of 

Gyno-dicecious,  where  the  flowers  on  separate  individuals  are 
some  hermaphrodite  and  some  female,  but  none  male  only ;  and 
Andro-dioecious,  of  hermaphrodite  flowers  and  male,  but  no 
separate  female.  The  latter  is  a  less  common  case. 

Neutral,  as  applied  to  a  flower,  denotes  that  both  stamens  and 
pistils  are  wanting,  —  a  case  neither  rare  nor  inexplicable  on 
grounds  of  utility.  (356,  504.) 

Sterile  and  Fertile  are  more  loosely  used  terms.  A  sterile 
flower  may  mean  one  which  fails  to  produce  seed,  as  a  sterile 
stamen  denotes  one  which  produces  no  good  pollen,  and  a 
sterile  pistil  one  which  is  incapable  of  seeding.  .  But  commonly  a 
sterile  flower  denotes  a  staminate  one  ;  a  fertile  flower,  one  which 
is  pistillate,  if  not  also  hermaphrodite. 

348.  Suppressed  Perianth.  Almost  universally,  when  the  peri- 
anth is  reduced  to  a  single  circle,  it  is  the  inner,  or  corolla,  which 
is  not  produced.  Or,  rather,  when  there  is  only  one  circle  or  sort 
of  perianth-leaves,  it  is  called  calyx,  whatever  be  the  appearance, 
texture,  or  color,  unless  it  can  somehow  be  shown  that  an  outer 
circle  is  suppressed.  For  since  the  calyx  is  frequently  delicate 
and  petal-like  (in  botanical  language,  petaloid  or  colored,  as  in 


192  THE  FLOWER. 

Clematis  and  Anemone,  Fig.  364) ,  and  the  corolla  is  sometimes 
greenish  or  leaf-like,  the  only  real  difference  between  the  two  is 
that  the  calyx  represents  the  outer 
and  the  corolla  the  inner  series. 
Even  this  distinction  becomes  arbi- 
trary when  the  perianth  consists  of 
three  or  four  circles,  or  of  a  less 
definite  number  of  spirally  arranged 
members. 

349.  Yet  the  only  perianth  obvi- 
ously present  may  be  corolla,  as  when 
the  calyx  has  its  tube  wholly  adnate 
to  the  ovary  and  its  border  or  lobes  obsolete  or  wanting.1  Aralia 
nudicaulis  (Fig.  341)  is  an  instance,  likewise  many  Umbelliferse, 
some  species  of  Fedia  or  Valerianella,  the  fertile  flowers  of  Nyssa, 
and  those  Compositae  which  have  no  pappus.  For  PAPPUS, 
the  name  originally  given  to  thistle-down  and  the  like,  answers 
to  the  border  or  lobes  of  a  calyx  attenuated  and  depauperated 
down  to  mere  fibres,  bristles,  or  hairs.  The  name  is  ex- 
tended to  other  and  less  obliterated  forms.  (644,  Fig.  631-633.) 
When  the  obliteration  is  complete,  as  in  Mayweed  (Fig.  630) , 
in  some  species  of  Coreopsis,  &c.,  the  corolla  seems  to  be  simply 
continuous  with  the  apex  of  the  ovary.  A  comparison  with 
related  forms  reveals  the  real  state  of  things.2 

350.  So  also  in  Hippuris,  in  which  (along 
with  extreme  numerical  reduction  of  the  other 
floral  circles)  the  calyx  as  well  as  corolla  seems 
to  be  wanting ;  but  the  insertion  of  the  stamen 
on  the  ovary  (epigynous)  suggests  an  adnate 
calyx,  and  near  inspection  detects  its  border. 

351.  Both  calyx  and  corolla  are  really  want- 
ing in  the  otherwise  complete   and  perfect  (symmetrical  and 

1  In  the  flowers  of  the  two  common  species  of  Prickly  Ash  (Zanthoxy- 
lum)  of  the  Atlantic  United  States,  one  has  a  double,  the  other  a  single 
perianth  (as  shown  in  Gray,  Gen.  Illustr.  ii.  148,  t.  156) :  the  position  of  the 
stamens  gives  a  presumption  that  the  missing  circle  of  the  latter  is  the  calyx ; 
yet  it  may  be  otherwise  explained.     In  Santalaceae  there  are  some  grounds 
for  suspecting  that  the  simple  perianth,  although  opposite  the  stamens,  is 
corolla ;  and  the  foliaceous  sepal-lobes  of  the  female  flowers  of  Buckleya 
would  confirm  this,  if  these  are  true  sepals  rather  than  adnate  bracts. 

2  In  the  pappus  of  Compositae,  every  gradation  is  seen  between  undoubted 
calyx,  recognizable  as  such  by  structure  as  well  as  position,  and  diaphanous 
scales,  bristles,  and  mere  hairs,  wholly  "  trichomes  "  as  to  structure,  although 
in  the  place  of  "  phyllomes  "   and  representing  them. 

PIG.  364.    Flower  of  Anemone  Pennsylvania ;  apetalous,  the  calyx  petaloH. 
FIG.  365.    Aclilamydeous  flower  of  Lizard'8-tail  (Saururus  cernuus),  magniied. 


DISAPPEARANCE   OF   PARTS. 


193 


trimerous)   flowers  of  Saururus,  Fig.  365.      But  achlamydeous 
blossoms  are  usually  still  further  reduced  to  a  single  sex. 

352.  Suppression  of  one  circle  of  stamens  is  of  very  common 
occurrence.  It  is  seen  in  different  species  of  Flax  ;  which  have 
mostly  5-merous  perfectly  symmetrical  and  complete  flowers  with 
one  set  of  stamens  abortive.  In  some  species  (as  in  Fig.  367), 


vestiges  of  the  missing  circle  of  stamens  are  conspicuous  in 
the  form  of  abortive  filaments,  interposed  between  the  perfect 
stamens ;  in  others,  these  rudiments  are  inconspicuous  or  even 
altogether  wanting. 

353.  Suppressed  Andrcecium  or  Gfynoecium.  This  occurs  with- 
out or  along  with  suppression  in  the  perianth.  In  cases  of  the 
former,  vestiges  of 
the  aborted  organs 
often  remain  to  sig- 
nify the  exact  nature 
of  the  loss.  Sepa- 
ration of  the  sexes 
(monoecious,  dioeci- 
ous, &c.)  is  the  re- 
sult of  such  suppres- 
sion. In  Menisper- 
mum  (Fig. 368,  369), 
this  is  accompanied 
by  an  actual  doub- 
ling of  both  calyx 
and  corolla.  The 
dioecious  flowers  of 
Smilax  are  similarly 
complete,  except  by 
the  abortion  of  one  sex,  but  the  calyx  and  corolla  are  single. 

FIG.  366.  Flower  of  a  Linum  or  Flax.  367.  Andrcecium  and  gynoacium ;  the  former 
of  5  perfect  stamens,  alternating  with  5  rudiments  of  a  second  set. 

FIG.  368,  369.  Dio3cious  flowers  of  Moonseed,  Menispermum  Canadense :  368,  Stami- 
nate  or  male  blossom ;  369,  Pistillate  or  female,  but  with  six  abortive  stamens,  before 
as  many  petals. 

FIG.  370.  A  catkin  of  staminate  flowers  of  a  Willow,  Salix  alba.  371.  A  single 
staminate  flower  detached  and  enlarged  (the  bract  turned  from  the  eye)  372  \  pistil- 
late catkin  of  the  same  species.  373.  A  detached  pistillate  flower,  magnified 


194 


THE  FLOWER. 


354.  Combined  with  suppression  of  Perianth.  This,  which  is 
found  in  most  amentaceous  or  catkin-bearing  trees,  in  some  with 
partial  suppression  of  perianth,  is  well  illustrated 
in  Willows,  the  flowers  of  which  are  all  achlamydeous 
and  dioecious.  (347.)  The  little  scale  (gland  or  nec- 
tary) at  the  inside  of  each  blossom  might  be  sup- 
posed to  represent  a  perianth,  reduced  to  a  single 
piece  ;  but  an  extended  comparison  of  forms  refers 
it  rather  to  the  receptacle.  Willow-blossoms  (Fig. 
370-372)  are  crowded  in  catkins,  each  one  in  the  axil  of  a  bract : 
the  staminate  flowers  consist  of  a  few  stamens  merely,  in  this 
species  of  only  two,  and  the  pistillate  of  a  pistil  merely.  In 
Salix  purpurea,  the  male  flower  seems  to  be  a  single  stamen 
(Fig.  374)  ;  but  it  consists  of  two  stamens,  united  into  one 

body.  Here  extreme 
suppression  is  ac- 
companied with  co- 
alescence of  the 
existing  members. 

355.  Still  more 
simplified  flowers, 
but  more  difficult 
to  comprehend,  are 
those  of  Euphorbia, 
or  Spurge.  These 
are  in  fact  monoeci- 
ous ;  and  the  female 
flower  is  a  pistil,  the 
male  is  a  stamen. 
The  pistillate  flower 
(of  three  carpels, 
their  ovaries  united 
into  one  three-lobed 

375  STB  compound      ovary) 

surmounts  a  slender  peduncle  which  terminates  each  branch  of  the 
flowering  plant.  (Fig.  375.)  From  around  the  base  of  this  pe- 
duncle rise  other  smaller  and  shorter  peduncles,  each  from  the 
axil  of  a  slender  bract,  and  surmounted  by  a  single  stamen, 
which  represents  a  male  flower.  (Fig.  376,  377.)  This  umbel-like 

FIG.  374.  A  separate  staminate  flower  of  Salix  purpurea,  with  the  stamens  coa- 
lescent  (monad elphous  and  syngenesious),  so  as  to  appear  like  a  single  one. 

FIG.  375.  Flowering  branch  of  Euphorbia  corollata.  376.  Calyx-like  involucre 
divided  lengthwise,  showing  the  staminate  flowers  around  a  pistillate  flower  (a).  377.  A 
more  magnified  staminate  flower  detached  with  its  bract,  a ;  its  peduncle  or  pedicel  b, 
surmounted  by  the  solitary  stamen,  c.  378.  Pistil  in  fruit,  cut  across,  showing  the 
three  one-seeded  carpels  of  which  it  is  composed. 


SUPERPOSITION  OF   SUCCESSIVE   PARTS.  195 

flower- cluster  is  surrounded  and  at  first  enclosed  by  an  involucre 
in  the  form  of  a  cup,  which  imitates  a  calyx  ;  and  the  lobes  of  this 
cup  (the  free  tips  of  the  calyx-leaves)  in  the  present  species  are 
bright  white,  so  that  they  exactly  imitate  petals.  Here,  then,  is 
a  whole  cluster  of  extremely  simplified  flowers,  taking  on  the 
guise  of  and  practically  behaving  like  a  single  flower,  the  invo- 
lucre serving  as  calyx  and  corolla  ;  the  one-stamened  male  flowers 
collective^  imitating  the  andrcecium  of  a  polyandrous  blossom, 
and  surrounding  a  female  flower  which  might  pass  for  the  pistil  of 
it.  A  series  of  related  forms,  from  various  parts  of  the  world, 
gives  proof  that  this  interpretation  is  the  true  one. 

356.  Suppression   of  both   Androecium   and  Gynoecium.      This 
occurs  in  what  are  termed  Neutral  Flowers  (347) ,  such  as  are 
conspicuous  at  the  margin  of  the  cymes  of  Hydrangea  (Fig.  293) 
and  of  Viburnum  lantanoides  and  Opulus,  also  at  the  margin  of 
the  head  of  flowers  of  Sunflower,  Coreopsis  (Fig.  287,  288),  and 
the  like.     In  these  and  most  other  instances,  the  perianth  of 
which  only  the  flower  consists  is  much  larger  and  more  showy 
than  in  the  accompanying  perfect  flowers :  in  fact,  their  whole 
utility  to  the  plant,  so  far  as  known,  is  in  this  conspicuousness. 
No  plant  normally  bears  neutral  flowers  only ;  but  in  cultivation 
all  sometimes  become  so  by  monstrosity,  as  in  the  form  of  Vibur 
num  Opulus  called  Snowball  or  Guelder  Rose,  also  in  "full 
double"  roses,  pinks,  &c.     Occasionally  flowers  become  sterile 
and  neutral  by  mere  depauperation  and  abortion  of  perianth  as  well 
as  of  essential  organs,   as  in  certain  Grasses ;   but  such  are 
mostly  vestiges  of  flowers  rather  than  neutral  blossoms. 

§  6.   INTERRUPTION  OF  NORMAL  ALTERNATION. 

357.  A nt ^position  or  Superposition  is  the  opposition  of  succes- 
sive (or  apparently  successive)  whorls  which  normally  alternate. 
This  result  is  brought  about  in  different  ways,  some  of  which  are 
obvious,  while  of  some  the  explanation  is  hypothetical. 

358.  In  the  first  place,  there  are  cases  of  seeming  anteposi- 
tion,  which  are  explained  away  on  inspection.     In  a  tulip,  lily, 
and  the  like,  there  is  a  perianth  of  six  leaves  and  a  stamen  be- 
fore each.     The   simple  explanation  is   that  the  flower  is  not 
6-merous,  but  3-merous  :  there  is  a  calyx  of  three  sepals,  colored 
and  mostly  shaped  like  the  three  petals,  which  alternate  with 
these  and  are  clearly  anterior  in  the  bud ;  next,  three  stamens 
alternate  with  the  petals  or  inner  circle  of  the  perianth ;  then 
the  three  stamens  of  the  inner  circle,  alternating  with  the  preced- 
ing, necessarily  are  opposite  the  three  petals,  as  the  first  three  are 


196  THE   FLOWER. 

opposite  the  sepals.  These  organs  altogether  are  in  four  whorls 
of  three,  not  in  two  of  six  members  ;  and  the  pistil  at  the  centre, 
of  three  combined  members,  is  the  fifth  and  final  whorl. 

359.  The  Barberry  family  exhibits  a  similar  seeming   ante- 
position,  which  is  more  striking  on  account  of  a  multiplication 
of  the  members  of  the  perianth.     The  calyx  is  of  six  sepals  in 
two  circles,  the  corolla  of  six  petals  in  two  circles,  the  stamens 
equally  six ;  and  so  each  petal  has  a  stamen  before  and  a  sepal 
behind  it.     But,  when  properly  viewed  as  a  trimerous  flower  with 
double  circles  of  sepals  and  petals  as  well  as  of  stamens,  all  is 
symmetrical  and  normal.    Menispermum  in  the  related  Moonseed 
family  is  in  the  same  case,  but  the  flower  is  trimerous,  as  seen 
in  Fig.  369  :  in  the  male  blossom  this  is  obscured  in  the  androe- 
cium   (Fig.    368)   by  a  multiph'cation  of  the   stamens.1     The 
same  thing  occurs  in  the  perianth  and  bracts  of  certain  Clusiaceae, 
in  which  the  members  counted  as  in  fours  are  superposed,  and 
in  some  of  which  the  double  dimerous  arrangement  with  apparent 
anteposition  extends  through  the  corolla  ;  while,  in  other  closely 
related  flowers,  the  corolla  changes  to  simply  tetramerous  and  to 
alternation  with  the  preceding  four  sepals.     This  passes,  in  the 
same  family  and  in  the  allied  Ternstroemiacese,  into 

360.  Superposition  by  Spirals,  as  where  five  petals  are  ante- 
posed  to  five  sepals,  by  an  evident  continuation  of  pentastichous 
phjilotaxy ;    and  the  stamen-clusters  of  Gordonia  Lasianthus 
are  probably  in  this  way  brought  before  the  petals.2     The  flower 
of  Camellia  is  continuously  on  the  spiral  plan  up  to  the  gynos- 
cium  ;  but  upon  one  which,  from  the  bracts  onward,  rises  from  the 
\  to  the  I  and  f  order  or  higher,  throwing  the  petals  of  the  rosette 
in  a  full-double  flower  into  numerous  more  or  less  conspicuous 
vertical  ranks. 

361.  Anteposition  in  the  Aiulnwiuni.     It  is  in  the  andrcecium 
that  real  anteposition  is  most  common,  and  also  most  difficult  to 
account  for  upon  any  one  principle.     Doubtless  it  comes  to  pass 
in  more  than  one  way.     This  condition  is  chiefly  noticed  when 
the  stamens  are  definite  in  number,  and  mainly  in  isostemonous 
and  diplostemonous  flowers.  (324.) 

362.  With  Isostemony.     Vitis  (Fig.  379-381),  also  Rhamnus 
(Fig.  415,  416),  and  the  whole  Grape  and  Buckthorn  families  of 

1  In  Columbine  (Aquilegia),  multiplication  of  the  stamens  in  successively 
alternating  5-merous  whorls  similarly  brings  the  andrcecium  into  ten  ranks ; 
so,  when  these  stamens  in  double  flowers  are  transformed  into  hollow-spurred 
petals,  these  are  set  one  into  another  in  ten  vertical  ranks. 

2  Gen.  Illustr.  ii.  1. 140.     But  the  petals  alternate  with  the  sepals  in  the 
ordinary  manner  of  the  flower,  though  their  strong  quincuncial  imbrication 
suggests  the  spiral  arrangement. 


SUPERPOSITION  OF   SUCCESSIVE   PABTS. 


197 


which  they  are  the  types,  afford  familiar  cases  of  a  single  circle 

of  stamens  placed  before  the  petals.     In  Vitis,  there  are  green 

nectariferous  lobes  or  processes  from 

the  receptacle,  alternate  with  and  inside 

the  stamens  :  there  is  no  good  reason  to 

suppose  that  they  answer  to  a  second 

row  of  stamens.    All  isostemonous  Por- 

tulacacese  have  the  stamens  before  the 

petals  ;  and,  when  the  stamens  are  fewer 

than  the  petals,  those  which  exist  occupy 

this  position.     Among  the  orders  with 

gamopetalous  corolla,  such  anteposition 

is  universal  in  Plumbaginacese,  Primu- 

laceae,  the  related  Myrsinacese,  and  in 

most  Sapotacese,  in  the  latter  usually 

with  some  complications. 

363.  The  earliest  and  the  most  obvious  explanation  of  the 
anomaly  is  that  of  the  suppression  of  an  outer  circle  of  stamens, 
and  to  this  view  recent  morphologists  are  returning.1  Observa- 
tion supplies  no  vestige  of  proof  of  it  in  Rhamnaceae  and  Vitaceae  ; 
but,  in  the  group  of  related  orders  to  which  the  Primulaceae  be- 
long, evidence  is  not  wanting.  For  Samolus  and  Steironema 
both  exhibit  a  series  of  rudimentary  organs  exactly  in  the  place 
of  the  wanting  circle  of  stamens,  which  may  well  be  sterile  fila- 
ments. In  the  allied  order  Sapotaceae,  while  Chrysophyllum 
has  in  these  respects  just  the  structure  of  Primulaceae,  and 
Sideroxylon  that  of  Samolus,  Isonandra  Gutta  (the  Gutta-percha 
plant)  has  a  circle  of  well-formed  stamens  in  place  of  the  sterile 
rudiments  of  the  preceding ;  that  is,  alternate  with  the  petals, 


1  Eichler,  Bliithendiagramme,  passim,  and  in  preface  to  Part  II.  xviii., 
relating  chiefly  to  obdiplostemony.  The  principal  opposing  view  is  that  of 
St.  Hilaire,  Duchartre,  &c.,  maintaining  that  corolla  and  stamens  here  repre- 
sent one  circle  of  organs  doubled  by  median  chorisis  ;  upon  which  see  note 
under  a  following  paragraph.  According  to  that  hypothesis,  there  is  no 
androecial  circle  in  such  blossoms,  or  only  vestiges  of  one,  but  the  petals  have 
supplied  the  deficiency  by  a  supernumerary  production  of  their  own !  The 
more  plausible  hypothesis  of  Braun,  that  of  a  suppressed  interior  circle 
of  extra  petals,  would  restore  the  alternation,  and  make  the  extant  sta- 
mens the  fourth  floral  circle,  as  does  the  adopted  explanation.  Braun's 
hypothesis,  if  it  insists  that  an  extra  row  of  petals  is  wanting,  supposes  the 
suppression  of  that  which  very  rarely  exists ;  but,  if  of  stamens,  then  the 
supposed  suppression  is  of  that  which  is  so  generally  present,  or  with  indi- 
cations of  presence,  as  properly  to  be  accounted  a  part  of  the  floral  type. 

FIG.  379.  Flower  of  the  Grape  Vine,  casting  its  petals  before  expansion.  380.  The 
same,  without  the  petals:  both  show  the  glands  of  the  disk  distinctly,  within  the 
stamens.  381.  Diagram  of  the  flower. 


198 


THE   FLO  WEB. 


completing  the  sjonmetry  of  the  blossom  and  the  normal  alterna- 
tion of  its  members.  This  explanation  of  the  anteposition  of  a 
single  circle  of  stamens  is  the  more  readily  received,  because  it 
well  accords  with  the  idea  here  adopted,  that  the  androscium 
of  a  typical  flower  should  consist  of  two  circles  of  stamens. 
(324.)  The  only  serious  objections  to  this  explanation  rise  out  of 
the  difficulty  of  applying  it  to  analogous  anteposition  when  both 
circles  are  present. 

364.  For  Diplostemony,  the  condition  of  two  circles  of  sta- 
mens, each  of  the  same  number  as  the  petals,  is  also  itself  very 
commonly   attended    by   anteposition.      In   normal    or   Direct 
Diplostemony,  —  that  which   answers   to   the   floral  type   com- 
pletely,—  the  antisepalous  stamens  (324,  note)  are  the  outer 
and  the  antipetalous  the  inner  series,   and  the  carpels  when 
isomerous  alternate  with  the  latter  and  oppose  the  sepals ;  the 
alternation  of  whorls  is  therefore  complete,  as  in  the  diagram, 
Fig.  382.     Such  stamens,  however,  may  actuall}-  occupy  a  single 

line  or  coalesce  into  a 
tube,  without  derange- 
ment of  the  type.  But 
it  as  commonly  occurs 
that  the  antipetalous  sta- 
mens are  more  or  less 
exterior  in  insertion,  and 
then  the  carpels,  when 
isomerous,  are  alternate 
with  the  inner  and  anti- 
sepalous stamens,  and  therefore  opposite  the  petals,  as  in  the 
diagram,  Fig.  383.  This  arrangement  takes  the  name  of 
Obdiplostemony.  In  it  the  normal  alternation  of  successive 
whorls  is  interrupted,  so  as  to  produce  anteposition, 

365.  With  Obdiplostemony.     This  condition  prevails,  more  or 
less  evidently,  in  Ericaceae,  Geraniaceae,  Zygophyllaceae,  Rutaceae, 
Saxifragaceae,  Crassulaceae,  Onagraceae,  &c.  (but  in  some  of  these 
with   exceptions   of  direct   diplostemony)  ;    also,   accompanied 
by  a  peculiar  multiplication  of  members  (380),  in  Malvaceae, 
Sterculiaceae,  and  Tiliaceae.     The  explanation  is  difficult.     The 
Irypotheses  may  be  reduced  to  three,  neither  of  which  is  quite 
satisfactory.     There  is,  first,  the  hypothesis  of  St.  Hilaire,  ap- 
plied to  this  as  to  the  preceding  case  (to  Rhamnus,  Vitis,  &c.), 
that  these  exterior  antipetalous  stamens  belong  to  the  corolline 
whorl ;  in  other  words,  that  the  petal  and  the  stamen  before  it 

.  FIG.  382.     Diagram  of  pattern  flower  with  direct  diplostemony,      383.   Diagram 
of  similar  flower  with  Obdiplostemony.    Both  from  Eichler'a  Bluthendiagramme. 


SUPERPOSITION   OF   SUCCESSIVE   PARTS.  199 

(whether  adnate  to  or  free  from  it)  answer  to  one  leaf  which 
has  developed  into  two  organs  by  a  deduplication  (372)  taking 
place  transversely.  This  makes  the  inner  and 
antisepalous  stamens  the  third  floral  circle 
or  the  only  truly  androecial  one,  and  sym- 
metrically alternate  with  the  petals  on  the  one 
hand  and  the  carpels  on  the  other.  The 
second  hypothesis  conceives  that  there  is  a 
whorl  suppressed  between  these  antipetalous 
stamens  and  the  corolla :  this,  ideally  restored, 
gives  symmetric  succession  and  alternation  to  all  the  succeeding 
whorls.  The  five  glands  in  a  Geranium-flower,  alternate  with  and 
next  succeeding  the  petals  (Fig.  384) ,  were  plausibly  supposed  to 
represent  this  missing  whorl,  which  according  to  Braun  should  be 
an  inner  corolla  ;  according  to  others  rather  a  primary  circle  of 
stamens.  The  third  is  the  recent  hypothesis  of  Celakowsky, 
which  Eichler  adopts :  this  regards  the  antipetalous  stamens  as 
really  the  inner  or  second  circle,  and  conceives  that  in  the  course 
of  development  it  has  become  external  by  displacement.  The 
difficulties  of  this  hypothesis  are,  first  to  account  for  this  dis- 
placement, and  then  for  the  anteposition  of  the  carpels  to  the 
assumed  inner  stamens  in  the  great  majority  of  these  cases.1 

1  In  the  first  part  of  the  Bliithendiagramme,  Eichler  inclined  to  the  first 
hypothesis,  that  of  St.  Hilaire  (now  very  much  abandoned  on  account  of 
the  feeble  evidence  that  there  is  any  such  thing  as  transverse  or  median 
chorisis);  in  the  second,  he  discards  this  in  favor  of  Celakowsky's  view 
(published  in  Regensburg  Flora,  1875).  As  to  members  which  are  morpho- 
logically interior  becoming  exterior  by  outward  displacement,  Eichler  cites 
the  staminodia  or  sterile  stamen-clusters  of  Parnassia  (Fig.  400,  401),  and 
the  corresponding  antipetalous  stamens  of  Limnanthes,  as  clearly  interior 
hi  the  early  flower-bud,  but  exterior  at  a  later  period  ;  states  that  the  vascu- 
lar bundles  which  enter  these  stamens  generally  are  either  inner  as  respects 
those  of  the  episepalous  stamens  or  in  line  with  them ;  that  in  some  cases 
(as  in  many  Caryophyllaceae)  the  real  insertion  of  the  stamens  is  that  of 
direct  diplostemony,  while  the  upper  part  of  their  filaments  and  the  anthers 
are  external  to  the  episepalous  series ;  that  in  most  families  with  obdiplos- 
temony  examples  of  direct  diplostemony  occur,  and  still  more  cases  with  both 
stamineal  circles  inserted  in  the  same  line ;  and  that,  as  a  rule,  the  episep- 
alous stamens  are  either  later  or  not  earlier  formed  than  the  epipetalous. 
As  to  the  position  of  the  carpels  before  antipetalous  stamens  and  petals, 
Celakowsky  suggests  that  this  may  result  from  the  outward  recession  of 
those  stamens  affording  more  room  there,  while  in  the  normal  case  the 
greater  space  is  over  the  episepalous  stamens.  And,  indeed,  exceptions 
to  the  prevalent  position  are  not  uncommon  both  in  direct  diplostemony 

FIG.  384.  Diagram  (cross- section)  of  the  flower  of  Geranium  maculatum,  exhibiting 
the  relative  position  of  parts,  and  the  symmetrical  alternation  of  circles,  ».  e.  sepals, 
petals,  greenish  bodies  called  glands,  antipetalous  stamens,  antisepalous  stamens, 
carpels. 


200  THE   FLOWER. 

366.  The  case  of  stamens  in  a  cluster  before  the  petals  is  a 
complication  of  either  of  the  foregoing  with  a  peculiar  kind  of 
multiplication,  termed  deduplication  or  chorisis.  (372.) 


§  7.     INCREASED  NUMBER  OF  PARTS. 

367.  Augmentation  in  the  number  of  floral  members  is   one 
of  the  commonest  modifications  of  the  type.     It  occurs  in  two 
ways  :  1st,  by  an  increased  number  of  circles  or  turns  of  spirals 
in  the  flower,  which  is  Regular  Multiplication ;  2d,  by  the  pro- 
duction of  two  or  three  or  of  many  organs  in  the  normal  place 
of  one,  Chorisis  or  Deduplication.     The  first  does  not  alter  the 
normal  symmetry  of  the  blossom,  although  it  may  render  it  dif- 
ficult or   impossible  to   trace  or  demonstrate  it.     The  second 
apparently  disturbs,    or  at  least    disguises,    floral   symmetry. 
Either  may  be   definite,  or  of  a  constant   and   comparatively 
small   number ;    or  indefinite,   when   too   numerous   for  ready 
counting,  or  inconstant,  as  the  higher  numbers  are  apt  to  be. 

368.  Regular  Multiplication,  or  Augmentation  of  floral  circles 
or  spirals,  may  affect  any  or  all  the  four  organs,  but  most  com- 
monly the  andrcecium.     When  the  perianth  is  much  increased 
in  the  number  of  its  members,  the  distinction  between  calyx  and 
corolla,  or  even  between  bracts  and  corolla,  is  apt  to  disappear, 
as  in  most  Cactaceous  flowers  (Fig.  317),  Nelumbium,  Calycan- 
thus,  &c.    In  these  and  similar  cases,  the  members  of  the  perianth 
are  prone  to  take  a  spiral  instead  of  cyclic  arrangement ;  and  this 


and  in  obdiplostemony.  Along  with  the  lack  of  clear  analogy  to  support 
St.  Hilaire's  hypothesis  of  transverse  deduplication,  the  similar  orientation 
of  the  vascular  bundles  in  the  petal  and  the  stamen  before  it  must,  as 
Celakowsky  insists,  be  good  evidence  that  these  represent  independent 
leaves,  and  not  superposed  portions  of  one. 

The  main  objection  to  the  second  hypothesis  (that  of  a  suppressed 
circle  outside  of  the  antipetalous  stamens)  is  that  this  missing  circle, 
whether  of  petals  or  stamens,  is  not  actually  met  with  in  any  nearly  re- 
lated forms  (for  in  Monsonia  the  fifteen  stamens  are  otherwise  explained) ; 
also  that  there  are  transitions,  as  above  mentioned,  between  obdiplostemony 
and  direct  diplostemony.  To  Braun's  theory  that  the  glands  behind  the 
antisepalous  stamens  in  true  Geraniaceae  answer  to  suppressed  phylla, 
Eichler  objects  that  these  are  present  behind  all  ten  stamens  in  Oxalideae ; 
also  that  all  are  wanting  when  the  office  of  nectar-secretion,  which  they  sub- 
serve, is  undertaken  by  some  other  part  of  the  flower,  as  by  the  calyx-spur 
in  Pelargonium  and  Tropaeolum.  The  first  objection  is  forcible  :  the  second 
mixes  morphological  considerations  with  functional,  and  is  inconclusive. 
Abortive  organs,  preserved  for  their  utility  as  nectaries,  might  totally  dis- 
appear when  rendered  useless  by  a  different  provision  for  the  same  function. 


INCREASED  NUMBER   OF   PARTS.  201 

is  even  more  true  of  greatly  multiplied  stamens  and  pistils,  as 
in  Magnolia  and  Liriodendron,  most  Anonaceae,  Ranunculus, 
Anemone,  and  the  like.  But  in  Aquilegia,  where  the  number  five 
is  fixed  in  the  perianth,  the  cyclic  arrangement  with  alternation 
of  whorls  prevails  throughout 

369.  The  definite  augmentation  of  calyx  and  corolla  by  the 
production  of  one  additional  whorl  of  each,  and  the  seeming 
anteposition  which  comes  of  it  when  the  andrcecium  remains 
simply  diplostemonous  (in   the   manner  of  the   Berberidaceae, 
Menispermaceae,  &c.,  359)  has  already  been  explained. 

370.  Similar  increase  to  two  whorls  affecting  the  corolla  only 
characterizes  Anonaceae,  Magnoliaceae,  Papaveraceae,  and  Fuma- 
riaceae.     In  all  but  the  last  order,  this  is  accompanied  by  indefi- 
nitely multiplied  stamens,  and  mostly  by  an  increased  number  of 
carpels.    In  Fumariaceae,  which  has  dimerous  flowers,  there  is  a 
diminution  by  the  suppression  in  most  cases  of  half  the  normal 
andrcecium,   and   also  an   augmentation   of  the   other  half  by 
chorisis.  (372.) 

371.  Parapetalous  Multiplication.     Under  this   head   may  be 
described   an   anomalous   arrangement  of  augmented  stamens 
which  prevails  in  the  order  Rosaceae,  but  is  not  pecuh'ar  to  it.1 
The  simplest  case,. but  a  rare  one,  is  seen  in  the  10-stamened  vari- 
ety of  some  Hawthorns,  as  occasionally  in  Crataegus  coccinea  and 
Crus-galli.     The  ten  are  in  one  circle  and  in  pairs,  the  pairs 
alternate  with  the  petals.     Some  would  say  the  pairs  are  before 
the  petals  ;  but  the  space  between  two  stamens  before  each  petal 
is  mostl}'  rather  wider  than  in  the  pair  taken  the  other  way. 
The  next  case  in  order,   as  in    15-stamened   Hawthorns,   and 
constantly  in  Nuttallia,  adds  to  the  above  a  simple  interior  circle 
of  five  stamens,  one  directly  before  the  middle  of  each  petal. 
Next,  as  in  most  Pomeae  and  many  PotentiUeae,  there  are  twenty 
stamens,  thus  placed,  but  with  an  additional  circle  of  five  alter- 
nating with  the  preceding  one.     Next  there  are  25  in  three 
circles,  the  second  circle  as  well  as  the  first  having  ten  stamens  ; 
and  finally  there  are  from  30  to  50,  all  probably  in  circles  of  ten 
each.     There  is  little  doubt  that  the  circles  develop  in  centri- 
petal order ;  the  inner  successively  the  later.2 

1  It  was  first  clearly  described  by  Dr.  A.  Dickson,  in  Trans.  Bot.  Soc. 
Edinb.  viii.  468,  and  Seemann's  Jour.  Bot.  iv.  473  (1866).   He  introduced  the 
term,  parapetalous,  which  is  characteristic  of  it  in  its  elementary  form  (254, 
note) :   it   is   particularly  illustrated  by  Eichler,  in  Bliithendiagramme,  ii. 
495-510.     The  former  interprets  it  by  chorisis,  both  median  and  collateral : 
the  latter  presents  the  facts  and  possible  views,  but  declines  to  adopt  either 
of  then). 

2  Accordingly,  the  whole  is  probably  to  be  explained  by  some  modifica- 

irTATE  TFT  • 
•&NTA  B>  'A 


202  THE   FLOWER. 

372.  Chorisis  or  Dednplication.     Both  these  terms,  and  the 
ideas  which  they  denote,  originated  with  Dunal,  but  were  first 
expounded  by  Moquin-Tandon.1     The  first  word  is  Greek  for  a 
separating  or  separation.     The  second  is  a  translation  of  Dunal's 
French  word  dedoubkment  (literally  undoubling) ,  the  ambiguity 
of  which,  and  of  the  original  presentation  of  the  case,   long 
retarded  the  right  apprehension  of  the  subject.     Diremption  has 
been  suggested  (by  St.  Hilaire)  as  a  proper  term.     The  mean- 
ing  simply  is,  the   division  of  that   which  is  morphologically 
one   organ  into  two  or  more   (a  division  which  is  of  course 
congenital) ,  so  that  two  or  more  organs  occupy  the  position  of 
one.     As  thus  used,  chorisis  is  restricted,  or  nearly  so,  to  the 
homologues  of  leaves  in  the  flower,  and  mainly  to  stamens  and 
carpels  ;  the  division  or  splitting  up  of  a  petal  or  a  sepal,  when  it 
occurs,  being  expressed  in  the  phrases  which  are  applied  to  leaves. 
Yet  a  compound  leaf,  especially  one  of  the  palmate  type,  is 
a  good  type  of  chorisis,  the  several  blades  of  a  compound  leaf 
answering  to  the  single  blade  of  a  simple  leaf.     It  has  been  ob- 
jected against  the  terms  chorisis  and  deduplication  that  they 
assume  the  division  of  that  which  has  never  been  united ;  but 
so  equally  does  the  established  terminology  of  foliage.     A  di- 
vided leaf  has  never  been  entire. 

373.  Chorisis  is  complete  when  the  parts  concerned  are  dis- 
tinct or  separate  to  the  very  insertion,  as  in  the  stamen-clusters  of 
Hypericum.     The  foliar  form  of  this  would  be  represented  by 

tion  of  the  augmentation  of  circles.  Dickson's  hypothesis,  that  the  two, 
three,  or  five  stamens  which  are  more  or  less  in  face  of  each  petal  are  all 
deduplications  of  that  petal,  would  come  to  be  noticed  under  the  next  head, 
but  it  may  be  dismissed  at  once.  Yet  that  the  pairs  in  the  outer  circle 
represent  each  an  antisepalous  stamen,  divided  by  chorisis  (sometimes 
incompletely)  and  much  separated,  is  not  improbable.  The  other  tenable 
explanation  (which  may  be  harmonized  with  the  last)  is  that  the  outer 
circle  of  stamens  here  rightly  consists  of  ten  members,  respectively  alternat- 
ing with  the  sepals  and  petals  taken  as  a  whole.  This  makes  them  para- 
petalous,  and  at  the  same  time  brings  them  under  Hofmeister's  general  law 
that  new  organs  originate  over  intervals  of  those  preceding,  in  this  case  over 
the  ten  perianth-intervals  directly.  It  also  accords\ith  Hartog's  elucidation 
of  the  accessory  parts  in  the  flower  of  Sapotaceae  (in  Trimen's  Jour.  Bot.  1878). 
The  inner  circles  are  there  sometimes  5-merous  after  the  primitive  type, 
sometimes  10-merous  in  regular  alternation  to  the  preceding  circles. 

1  Moquin-Tandon,  Essai  des  Dedoublemens,  &c.,  Montpellier,  1826 ;  Con- 
side'rations  sur  les  Irre'gularite's  de  la  Corolle,  &c.,  in  Ann.  Sci.  Nat.  xxvii. 
237,  1832 ;  Teratologie  Ve'getale,  337.  Dunal,  Essai  sur  les  Vaccinie'es, 
1819,  cited  by  Moquin  (some  pages  printed,  but  never  published) ;  Conside"- 
rations  sur  la  Nature  et  les  Rapports  de  quelques-uns  des  Organes  de  la 
Fleur,  1829.  The  next  botanist  to  develop  it  was  St.  Hilaire,  Morphologic 
Vege-tale,  1841. 


CHOEISIS   OR   DEDUPLICATION.  203 

such  sessile  palmately  compound  leaves  as  those  of  some  species 
of  Aspalathus.  It  is  incomplete  when  division  does  not  extend 
to  the  base ;  as  in  Fig.  387,  393.  .  Compare,  as  a 
proximate  homologue  of  this,  a  petal  of  Mignonette, 
Fig.  385.  But  proper  chorisis  requires  that  the 
supernumerary  organs  should  be  developed  like 
unto  the  original  organ  which  is  thus  multiplied,  or 
should  complete  their  symmetry,  whatever  it  be. 

374.  St.  Hilaire  distinguished  two  kinds  of  deduplication ;  viz., 
collateral  when  the  members  stand  side  by  side,   and  parallel 
when  an  organ  becomes  double  or  multiple  antero-posteriorly. 
The  latter,  sometimes  called  vertical,  and  sometimes  transverse, 
is  better  named  median  chorisis.     The  collateral  is  the  origi- 
nal and  typical  chorisis.    Most  botanists  incline  to  restrict  the 
name  to  this,  and  to  give  some  other  explanation  and  name  to 
the  median  form  of  augmentation.     But  some  cases,  such  as 
those  of  Tilia  and  Sparmannia,  are  clearly  of  the  same  nature 
as  the  collateral,  and  may  be  a  disguised  form  of  it ;  there  are 
others  which  may  be  explained  in  accordance  with  it ;  and  there 
are  such  transitions  between  some  of  these  and  coronal  out- 
growths that  the  term  chorisis  is  most  conveniently  made  to 
comprise  augmentation  or  doubling   in  either  plane.     Distinct 
anteposition,  however,  may  be  explained  in  other  ways.  (357.) 

375.  Typical  or  Collateral  Chorisis,  in  which  the  members, 
together  answering  to  one  leaf,  normally  stand  side  by  side, 
occurs  in  many  families  of  plants,  and 

in  a  variety  of  forms.    A  few  are  here 
presented. 

376.  Elodes  Virginica  (a  common 
marsh  plant  of  the  tlypericum  family) , 
like  most  of  its  near  relatives,  has  its 
calyx  and  corolla  on  the  plan  of  five, 
its  stamens  and  carpels  on  the  plan  of 

three,  as  is  shown  in  the  diagram,  Fig.  386.  This  makes  a  break 
in  the  symmetry  between  the  corolla  and  the  stamens ;  but  all 
within  is  in  regular  alternation  when  the  three  stamens  of  each 
cluster  are  counted  as  one  as  their  union  at  base  into  a  phalanx 
(Fig.  387)  may  suggest.  These  phalanges  alternate  with  the 
three  carpels,  and  therefore  stand  where  single  stamens  belong. 
The  three  conspicuous  green  projections,  which  in  a  general  way 

PIG.  385.  A  petal  of  Mignonette  (Reseda  odorata),  with  many  parted  blade, 
enlarged. 

FIG.  386.  Diagram  of  flower  of  Elodes  Virginica,  with  three  phalanges  of  stamens 
forming  the  inner  circle,  and  three  glands  answering  to  the  outer  circle.  387.  A  de- 
tached phalanx  of  three  stamens. 


204 


THE   FLOWER. 


are  called  glands,  alternate  with  the  phalanges,  and  so  are  taken 
to  represent  the  outer  circle  of  stamens.  The  morphologist 
accordingly  sees  in  the  glands  the  homologues  or  representatives 
of  the  outer  series  of  stamens,  reduced  to  three  by  abortion,  and 
in  the  three  stamen-clusters  only  the  three  alternating  stamens 
of  the  inner  series,  trebled  by  chorisis,  and  this  chorisis  incom- 
plete, because  it  has  not  quite  divided  the  filament  into  three. 
In  Hypericum,  the  glands  are  completely  suppressed,  each  pha- 
lanx is  almost  or  quite  divided  into  a  cluster,  either  of  about 
three  stamens  each,  as  in  H.  Sarothra,  or  of  a  few  more  (in  H. 
mutilum  and  H.  Canadense),  or  of  an  indefinite  number,  as  in 
the  common  St.  Johnsworts.  Then  in  some  other  species  (as 
in  our  H.  pjTamidatum)  the  carpels  and  the  stamen-clusters  rise 
to  five,  realizing  complete  pentamerous  symmetry,  except  that 
the  almost  numberless  stamens  all  belong  to  the  one  inner  circle. 
Morphologically,  they  are  comparable  to  the  leaflets  of  five  (or 
in  most  species  three) 
decompound  and  ses- 
sile or  almost  sessile 
leaves.  The  indefinitely 
numerous  stamens  of 
Ricinus  are  similarly 
increased  from  five  by 
compound  ramification. 
377.  Fumariacese,  the 
Fumitory  family,  may 
furnish  the  next  illus- 
tration. The  flower  is 
on  the  plan  of  two 
(dimerous)  throughout. 


Taking  Dicentra  to  show  it,  there  is  first  a  pair  of  small  and 
scale-shaped  sepals,  not  unlike  the  pair  of  bractlets  on  the 

PIG.  388.  Dicentra  Cucullaria  (Dutchman's  Breeches),  a  scape  in  flower  and  a  leaf, 
severed  from  the  singular  bulb  (formed  of  the  enlarged  bases  of  petioles).  389.  Detached 
flower,  of  natural  size,  showing  also  the  pair  of  bractlets  on  the  pedicel.  390.  Same 
with  parts  displayed,  and  391,  inner  petals  placed  above.  392.  Diagram  of  flower  of 
Dicentra  or  Adlumia,  from  a  section  across  the  summit.  393.  One  of  the  phalanges  of 
stamens  of  Adlumia ;  upper  part  only- , 


CHOEISIS   OK   DEDUPLICATION.  205 

pedicel  below  (Fig.  389,  390) ,  and  normally  alternate  with  them : 
alternate  with  these  is  a  pair  of  large  petals,  deeply  saccate,  or 
spurred  below ;  alternate  with  these,  a  pair  of  smaller  petals 
with  spoon-shaped  tips  which  cohere  at  the  apex  (the  corolla 
therefore  of  two  circles  as  in  the  related  Poppy  family)  ;  alternate 
with  these,  two  phalanges  or  united  stamen-clusters,  of  three 
stamens  each ;  alternate  with  these  is  nothing,  for  the  second 
set  of  stamens  is  wanting  ;  alternate  with  this  vacancy  is  a  pair 
of  carpels  wholly  combined  into  a  compound  2-merous  pistil. 
The  statement  itself  explains  the  morphology.  The  three  sta- 
mens of  each  phalanx  stand  in  the  place  of  a  stamen,  and  are 
the  divisions  of  one.  In  Dicentra  the  members  of  the  phalanx 
are  almost  separate ;  in  Adlumia  (Fig.  393)  and  Corydalis  the 
undivided  filament  reaches  almost  up  to  the  anthers.  The  middle 
anther  of  the  phalanx  is  normal,  or  two-celled ;  the  lateral 
anthers  are  one-celled,  as  if  halved.1 

1  Eichler  adopts  this  interpretation  (proposed  in  Gray,  Gen.  Illustr. 
i.  118),  and  applies  it  to  the  crucial  instance  of  Hypecoura.  In  the  flower 
of  this  Old  World  genus,  there  are  four  apparently  simple  and  complete 
stamens,  one  before  each  petal :  the  simplest  interpretation  would  be  that 
which  the  facts  appear  to  present,  viz.  that  both  dimerous  circles  of  stamens 
are  complete  and  normal.  But  Eichler  —  in  view  of  the  early  development 
and  the  double  vascular  bundles  of  the  stamens  before  the  inner  petals,  and 
some  occasional  slight  disjunction  of  their  anther-cells  —  considers  that 
the  interior  stamen-circle  is  wanting  here,  no  less  than  in  the  other  genera 
of  the  order ;  that  what  here  takes  its  place  before  each  inner  petal  is  a 
stamen  composed  of  the  adjacent  lateral  member  of  the  phalanx,  congeni- 
tally  severed  from  the  group  to  which  it  belongs  and  soldered  into  one  fila- 
ment, bearing  the  two  one-celled  anthers  so  brought  together  as  to  imitate  a 
normal  two-celled  anther.  The  organogeny  of  the  blossom  is  thought  to 
favor  this  hypothesis ;  and  it  certainly  favors  the  view  here  adopted  of 
the  composition  of  the  three-membered  phalanx  of  the  family  generally. 
If  this  interpretation  of  Hypecoum  seems  far-fetched,  it  is  no  more  so  than 
its  exact  counterpart,  through  which  DeCandolle,  Lindley,  and  others  explain 
the  case  of  the  rest  of  the  family.  Starting  with  that  genus  as  the  simple 
type,  they  conceive  that  the  stamen  opposed  to  each  inner  petal  is  each 
severed  into  two,  and  that  these  half -stamens  attached  to  the  sides  of  the  two 
intact  stamens,  thus  producing  the  phalanges  by  coalescence. 

A  good  empirical  conception  of  the  formation,  from  a  single  leaf,  of  three 
stamens  in  Fumariacese,  or  two  in  Cruciferae,  is  afforded  by  the  petals  of 
Hypecoum,  as  illustrated  by  Eichler.  The  outer  petals  are  slightly  three- 
lobed  from  the  apex;  the  inner  are  deeply  so  and  narrower.  The  mem- 
bers of  the  next  circle  in  the  family  generally  are  just  such  three-lobed 
bodies,  the  tip  of  each  lobe  transformed  into  an  anther.  There  is  an  ap- 
parent congruity  in  the  production  by  the  symmetrical  middle  lobe  of  a 
symmetrical  two-celled  anther,  and  of  a  one-celled  anther  by  each  unsym- 
metrical  lateral  lobe  or  stipule-like  portion.  A  fuller  development  of  these 
sides  of  the  leaf,  and  non-development  of  the  middle  portion  (somewhat 
after  the  analogy  of  Lathyrus  Aphaca,  Fig.  219),  with  anther-formation, 
would  convert  the  leaf  into  a  pair  of  stamens 


206 


THE   FLOWER. 


378.  The  obvious  relationship  of  Cruciferae  to  Fumariaceae, 
their  agreement  in  the  rare  peculiarity  of  having  the  two  carpels 
395  side  by  side  instead  of  fore  and  aft 

(median),  and  the  characteristic 
anomaly  which  the  androecium  pre- 
sents (z.  e.  the  tetradynamy ) ,  would 
give  reason  to  expect  that  its  prob- 
lems might  be  solved  by  chorisis. 
Indeed,  the  doctrine  was  applied  to 
this,  long  before  its  application  to 
the  other  order.  Beginning  at  the 
centre  (Fig.  395,  &c.),  the  pistil  is 
of  two  carpels,  right  and  left ;  alter- 
nate with  these  is  a  pair  of  stamens 
on  the  side  next  the  axis,  matched 
by  another  pair  on  the  opposite 
side  of  the  pistil,  the  four  longer 
and  interior  stamens ;  alternate 
with  these,  and  lower  in  insertion^ 
a  single  stamen  on  each  side  ;  next, 
lour  petals,  of  somewhat  various  overlapping  in  aestivation, 
which  essentially  alternate  with  the  two  single  stamens  and  the 
cwo  pairs  ;  lastly,  four  sepals,  alternating  with  the  four  petals  as 
a  whole,  the  anterior  and  posterior  overlapping  the  lateral  ones 
in  the  bud.  Now  the  median  (i.  e.  the  anterior  and  posterior) 
pairs  of  stamens  occasionally  have  their  contiguous  filaments 
conjoined,  as  in  Fig.  397.  If  this  were  at  all  constant,  the 
inference  would  undoubtedly  be  that  the  case  is  one  of  chorisis, 
and  that  the  flower  as  to  its  essential  organs  is  dimerous.  This 
is  apparently  the  best  explanation  to  be  given.  It  assumes  that 
the  chorisis  is  normally  complete  in  the  androecium  of  Cruciferae, 
instead  of  incomplete,  as  in  Fumariaceae.1  And  this  view  is 
confirmed  by  the  fact  that  the  median  stamens  are  simple  and 


1  The  hypothesis  here  adopted,  as  to  the  androecium,  is  that  of  Steinheil 
(1839),  and  of  Eichler  (in  Flora,  1865,  1872,  and  Bliithend.  ii.  200),  replacing 
that  of  Kunth,  1833,  &c.,  employed  in  former  editions.  The  rejected  view 
makes  the  flower  4-merous  up  to  the  pistil,  and  the  stamens  all  of  one  circle, 
alternating  with  the  four  petals,  the  median  stamens  (as  in  our  view)  doubled 
by  chorisis.  Krause  and  Wretschko  (cited  as  above  by  Eichler)  would 
have  the  floral  circles  2-merous  and  4-merous  by  turns ;  the  calyx  of  two 
2-merous  circles  (which  it  plainly  is) ;  the  corolla  of  one  4-merous  circle 

FIG.  394.  A  cruciferous  flower.  395.  Diagram  of  such  a  flower,  with  position  of 
axis  marked  above  it.  396.  Tetradynamous  stamens  and  the  pistil.  397.  A  common 
monstrosity  of  the  same,  two  of  the  four  inner  stamens  combined  into  a  common 
2-antheriferou8  body. 


CHORISIS   OK   DEDUPUCATION.  207 

single  in  Senebiera  and  many  species  of  Lepidium,  in  which  the 
lateral  or  short  stamens  are  at  the  same  time  abortive. 

379.  It  is  quite  possible  that  chorisis  may  be  extended  to  the 
corolla  of  the  cruciferous  flower,   and  reduce  the  whole  to  a 
symmetrical  2-merous  plan,  and  to  congruity  in  the  perianth 
also  with  Fumariaceae.     The  only  obstacle  is  in  the  petals  form- 
ing a  whorl  of  four  where  all  the  rest  is  2-merous,  for  the  sepals 
are  manifestly  two  decussating  pairs.     Now  the  median  petals 
of  Hypecoum  are  deeply  3-lobed.     An  abortion  of  their  middle 
lobe  would  leave  them  almost  two-parted :  a  little  more  would 
separate  them  ;  then  they  would  imitate  the  four  cruciferous  petals 
as  in  the  diagram,  Fig.  395.     Applying  this  view  to  Cruciferae, 
the  blossom  in  the  two  orders  would  accord  in  having  a  2-merous 
three- whorled  perianth,  the  first  and  third  whorls  median ; *  as 
also  in  the  dimerous  androecium,   the  first  whorl  of  which  is 
lateral.     The  difference  is  that  in  Fumariacese  the  two  members 
of  the  first  whorl  of  stamens  augment  by  chorisis  into  three,  and 
the  second  is  wanting,  or  is  present  only  in  Hypecoum  ;  while  in 
Cruciferae  the  first  whorl  is  simple  (of  the  two  short  stamens) ,  and 
the  second  is  doubled.    In  Fumariacese  only  the  first  whorl  of  the 
perianth  counts  as  calyx,  and 

the  corolla  is  of  two  whorls  ; 
in  Cruciferae,  the  first  and 
second  whorls  are  calyx,  the 
inner  sepals  answering  to  the 
outer  petals  of  Fumariaceae. 

380.  Chorisis    along  with 
anteposition  of  stamens  is  well 

seen  in  Tilia  or  Linden,  at  least  in  the  American  species.  In 
these  the  indefinitely  numerous  stamens  are  in  five  clusters,  one 
before  each  petal  (Fig.  398,  399),  and  there  is  a  petal-like  body 


alternating  with  the  calyx-members  as  a  whole ;  the  short  stamens  following 
as  a  2-merous  circle ;  then  the  long  stamens  as  a  4-merous  circle ;  lastly  the 
2-merous  gyncecium.  G.  Henslow  (in  Trans.  Linn.  Soc.  ser.  2,  i.  195)  would 
have  the  flower  4-merous  by  the  suppression  of  the  fifth  members  of  a 
6-merous  type,  and  a  further  suppression  of  half  of  the  remaining  exterior 
stamen-circle,  &c.  Finally,  there  is  the  much  better-maintained  view  that 
the  cruciferous  flower  is  2-merous  throughout,  as  explained  in  the  following 
paragraph,  379. 

1  This  view  was  taken  by  Steinheil,  in  Ann.  Sci.  Nat.  ser.  2,  337  (1839), 
and  is  essentially  reproduced  by  a  Russian  botanist,  Meschajeff,  in  Bull. 
Soc.  Imp.  Nat.  Mosc.  1872. 

FIG.  398.  Diagram  of  the  flower  of  Tilia  Americana,  the  common  American  Lin- 
den or  Basswood. 

FIG.  399.    A  detached  stamen-cluster  with  its  petal-like  scale. 


THE   FLOWER. 


in  each  cluster  with  which  the  stamens  cohere.  The  explanation 
by  chorisis  is  that  each  cluster,  petal-like  body  included,  is  a 
multiplication  of  one  stamen.  The  diagram  (Fig.  398)  accu- 
rately shows  that  most  of  the  stamens  originate  from  the  outer 
side  of  the  base  of  the  petal-like  portion :  this  is  most  naturally 
explained  by  median  chorisis.  The  superposition  of  the  clusters 
to  the  petals  will  take  the  same  explanation  as  that  of  Rhamnus, 
Vitis,  &c.  (Fig.  363.)  That  the  androecium  is  here  composed 
of  the  inner  circle  merely  is  partly  confirmed  by  the  alternation 
of  the  carpels  with  the  clusters.  According  to  Duchartre,1  the 
development  of  the  androecium  in  a  Mallow  indicates  a  similar 
structure  ;  for  the  whole  united  mass  originates  from  five  protu- 
berances, one  before  each  forming  petal  and  connected  with  it, 
this  by  collateral  chorisis  forming  a  cluster  of  stamens,  and  the 
five  clusters  coalescing  as  they  develop  into  a  tube  of  filaments, 
such  as  in  Fig.  485.  Now  Hibiscus  and  its  near  relatives  have 
a  naked  tip  to  the  stamen-tube,  ending  usually  in  five  teeth ; 
and  Sidalcea,  as  is  most  strikingly  shown  in  the  Californian 
S.  diploscypha,  has  two  series  of  stamens,  the  outer  (answering 
to  those  of  Malva  and  its  relatives)  in  five  membranaceous  pha- 
langes, superposed  to  the  petals ;  the  rather  numerous  inner 
series,  more  or  less  in  phalanges,  surmounts  an  interior  filament- 
tube.  Whence  it  is  inferred 
that  these,  and  the  five  teeth 
terminating  the  column  in 
Hibiscus,  represent  the  in- 
ner stamineal  circle  which  is 
wanting  in  Malva,  as  it  is  in 
Tilia.2 

381.  The  case  of  Paraas- 
sia  would  be  explained  as 
analogous  to  that  of  Tilia, 
but  with  the  stamen-clusters  before  the  petals  wholly  sterile, 
and  of  fewer  divisions,  while  an  inner  circle  of  five  stamens 

1  Comptes  Rendus,  1844,  &  Ann.  Sci.  Nat.  ser.  3,  iv.  123.    Duchartre  and 
others  who  draw  freely  upon  median  chorisis  to  explain  anteposition,  and 
consider  that  congenital  union  proves  it,  take  the  phalanges  in  these  cases, 
like  the  single  stamens  in  Vitis,  to  be  an  inner  part  of  the  petal  itself.    But 
this  view  appears  to  have  had  its  day. 

2  Gray,  Gen.  Illustr.  ii.  44,  57,  75-82.     The  position  of  the  carpels  before 
the  petals  in  Pavonia  and  Malvaviscus  brings  the  former  into  symmetrical 
alternation  with  such  an  inner  stamen-circle ;  but  it  is  not  so  in  Hibiscus, 
which  has  the  carpels  before  the  sepals. 

FIG.  400.    A  petal  of  Parnassia  Caroliniana,  with  a  triple  gtaminodium  before  it. 
FIG.  401.    Diagram  of  the  flower  of  Parnassia  Caroliniana. 


OUTGROWTHS.  209 

alternate  with  the  petals  forms  the  effective  androecium.  For 
the  scale-like  body  before  each  petal,  and  even  slightly  adnate 
to  its  base  (in  P.  Caroliniana  about  3-parted,  as  in  Fig.  400,  but 
in  P.  palustris  a  thin  scale,  fringed  with  more  numerous  gland- 
tipped  filaments) ,  is  plainly  outside  the  stamens  in  the  full-grown 
flower-bud.  But  Eichler  and  Drude  have  found 
that  it  is  inside  in  the  early  bud.1  Wherefore,  if 
these  stamen-like  bodies  really  represent  a  circle 
of  the  andro2cium,  it  must  be  the  inner  one  ;  and 
that  is  the  more  probable  view. 

382.  Multiplication  by  chorisis  in  the  gyncecium 
is  not  common ;   but  there  are  well  marked  in- 
stances of  it  in  all  degrees.     In  Drosera,  the 
styles   and  stigmas  are  doubled  (Fig.  402)  ;   in 
Malvaceae,  the  same  thing  takes  place  in  Pavonia 
and  its   allies;  while  in  Malope  and  two  other 

genera  of  the  same  order  the  few  normal  carpels  are  multiplied, 
evidently  by  chorisis,  into  an  indefinite  number  of  wholly  distinct 
ones. 

§  8.   OUTGROWTHS. 

383.  Proper  chorisis  is  the  congenital  multiplication  of  one 
organ  into  two  or  more  of  the  same  nature  and  office ;  or  at 
least  into  two  or  more  organs,  even  if  dissimilar,  as  in  the 
American  Lindens,  in  which  one  member  of  the  cluster  is  a  kind 
of  petal.     Between  this  and  the  production  by  an  organ  of  ap- 
pendages, or  outgrowths  of  little  or  no  morphological  signifi- 
cation, there  are  many  gradations ;  as  also  between  these  and 
mere   cellular    outgrowths   from    the   surface,    even    down    to 
bristles  and  hairs.     The  latter,  in  all  their  variety  and  modifica- 
tions, are  properly  outgrowths  of  the  epidermis  only,  and  there- 
fore consist  of  extended  cells,  single  or  combined,  unaccompanied 
by  vascular  or  woody  tissue.     To  them  has  been  given  the 
general  name  of  Trichomes  (Trichoma,  pi.  trichomata),  that  is 
structures  of  which  hairs  are  the  type.     They  may  occur  upon 
the  surface  of  any  organ  whatever.     Their  morphology  is  the 
morphology  of  cells  rather  than  of  organs.     They  will  therefore 
be  most   conveniently  illustrated  under  Vegetable  Anatomy  as 

1  Eichler  in  Fl.  Brasil.,  Sauvagesiacae,  &  Bliithend.  ii.  424 ;  Drude  in  Lin- 
naea,  xxxix.  239.  Eichler  refers  to  this  as  a  confirmation  of  Celakowsky's 
explanation  of  obdiplostemony  by  posterior  displacement.  (365.) 

FIG.  402.    Pistil  of  Drosera  filiformia  with  tricarpellary  ovary  (transversely  divided), 
and  six  styles,  i.  e.  three,  and  each  two-carted. 


210  THE  FLOWER. 

respects  their  structure,  and  in  the  Glossary  as  respects  ter- 
minology. 

384.  But  into  some  bristles,  such  as  those  of  Drosera,  a  sub- 
jacent stratum  of  tissue  enters,  including  one  or  more  ducts  or 
even  some  woody  tissue.     Prickles  are  of  this  class  ;  and  from 
the  most  slender,  which  pass  into  bristles,  there  are  all  grada- 
tions of  stoutness  and  induration.    Such  outgrowths  may  even  be 
formed  in  most  regular  order,  as  the  prickles  on  the  calyx-tube 
of  Agrimonia  and  scales  on  the  acorn-cup  of  Oaks,  and  yet  have 
no  morphological  importance.    On  the  other  hand,  true  represen- 
tatives of  leaf  or  stem  may,  by  abortion  and  depauperation,  be 
reduced  to  the  structure  as  well  as  the  appearance  of  trichomes. 
Examples  of  this  are  familiar  in  the  pappus  (answering  to  limb 
of  the  calyx)  of  many  Composite,  and  in  the  bristles  which 
answer  to  perianth  in  many  Cyperaceae.     The  scarious  stipules 
of  Paronychia  and  of  Potamogeton,  the  ligule  of  Grasses,  and 
even  the  corolla  in  Plantago,  are  equally  reduced  to  mere  cellular 
tissue.     So  that  the  structural  difference  between  trichomes  and 
outgrowths 1  is  not  at  all  absolute,  and  the  morphological  distinc- 
tion must  rest  upon  other  ground  than  anatomical  structure. 

385.  Among  the  corolline  outgrowths  most  akin  to  chorisis  is 
the  Crown  (  Corona)  of  Silene  and  allied  Caryophyllacese,  at  the 

junction  of  the  claw 
with  the  blade  of 
the  petals  (Fig.  403), 
the  analogy  and 
probable  homology 
of  which  to  the  ligule 
of  Grasses  (Fig. 
150)  is  evident;  also 
the  many-ra3'ed  fila- 
404  *»  mentous  crown  of 

Passion-flowers  (Fig.  404) ,  which  consists  of  two  or  more  series 
of  such  outgrowths.  In  Sapindus  and  some  other  Sapindacese, 
these  ligular  outgrowths  or  internal  appendages  are  more  like 
a  doubling  of  the  petal ;  as  also  in  Erythroxylum,  where  they 

*  This  is  the  best  English  name  for  the  Emergenzen  of  the  Germans,  the 
Epiblastema  of  Warming,  &c.  For  the  development  and  discussion  of  this 
subject,  see  Warming,  in  Kjobenhavn  Vidensk.  Meddel.  1872,  and  a  larger 
treatise  on  Ramification  in  Phanerogams,  Copenhagen,  1872.  Also,  Uhl- 
worm  in  Bot.  Zeit.  1873;  Celakowsky  in  Flora,  1874;  and  Eichler's  note  on 
Emergenzen  in  Bliithendiagramme,  i.  48. 

FIG.  403.    Petal  of  Silene  Pennsylvanica,  with  its  crown. 
FIG.  404.    Flower  of  Passiflora  cuerulea,  reduced  in  size. 


FORMS   OP  THE   RECEPTACLE.  211 

are  often  more  complicated  in  structure.     They  are  always  on 
the  inner  face,  and  are  commonly  two-lobed  or  parted. 

386.  Similar  stamineal  appendages  are  well  known  in  Cuscuta 
(Dodder) ,  in  Larrea  (Fig.  405)  and  other  Zygophyl- 

lacese,  and  less  conspicuously  in  Gaura. 

387.  To  extend  to  them  the  name  of  LIGTJLE  may 
not  be  amiss,  whether  they  are  regarded  as  mere 
outgrowths  of  floral  leaves,  without  further  morpho- 
logical relations,  or  whether  they  be,  at  least  some- 
times, interpreted  as  the  homologue  of  intrapetiolar 
stipules,  as  their  ordinarily  two-cleft  form,  and  their        m 
coincidence  in  Erythroxylum  with  an  intrapetiolar  two-cleft  stipule 


§  9.     FORMS  OF  THE  TORUS  OR  RECEPTACLE. 

388.  Torus  is  the  more  specific  and  proper  name,  RECEPTACLE 
is  the  more  usual.    (303.)     A  normal  receptacle  of  the  flower 
would  be  that  of  Fig.  316,  the  apex  of  the  flower-stalk  somewhat 
enlarged,  roundish  or  depressed,  and  with  surface  mainly  cov- 
ered by  the  insertion  of  the  several  organs ;  the  several  inter- 
nodes    which   it    potentially  contains   being 
undeveloped.     As  the  members  of  the  flower 

multiply  and  occupy  numerous  ranks,  the 
receptacle  enlarges  or  lengthens  to  give  them 
insertion  or  standing-room. 

389.  Of   elongated    forms   of   receptacle, 
Magnolia  and  Liriodendron  or  Tulip-tree  give 
familiar  instances.     The  lengthening  in  the 
former  is  mainly  for  the  support  of  both  an- 
droecium  and  gynoecium ;  in  the  latter,  as  in 
Myosurus,  mainly  for  the    gyncecium  only. 
The  fall  of  the  matured  carpels  reveals  it 
as   a  very   slender   or    bodkin-shaped   pro- 
longed axis.    Of  broadened  forms,  the  Straw- 
berry, even  in  blossom,   affords   a   familiar 
example.   (Fig.  406.)     In  the   same  order, 
Rubus  odoratus  shows  a  very  broad  and  flat 

receptacle :  in  roses,  it  is  so  deeply  concave  as  to  become  the 
reverse  of  the  strawberry  (Fig.  407),  being  urn- shaped  with  a 
narrow  mouth,  upon,  which  the  petals  and  stamens  are  borne, 

FIG.  405.  Stamen  of  Larrea  Mexicana,  with  a  conspicuous  ligulate  appendage  at 
the  base  within. 

FIG.  406.  Receptacle  of  a  strawberry  in  longitudinal  section.  407.  Same  of  a  row, 
in  diagram. 


212  THE  FLOWER. 

while  the  pistils  line  the  walls  of  the  cavity,  the  base  or  centre 
of  this  cavity  answering  to  the  apex  of  the  strawberry. 

390.  Sometimes  internodes  are  lengthened  between  certain 
members.     In  Schizandra,  the  receptacle,  barely  oblong  in  blos- 
som, lengthens  greatly  in  fruiting,  so  as  to  scatter  the  carpels 
on  a  long  filiform  axis. 

391.  In   many   Gentians,   in    Stanleya   and   Warea   among 
Cruciferse,  and  in  most  species  of  Cleome,  the  internode  of  the 

receptacle  between  stamens 
and  pistil  is  developed  into 
a  long  stalk  to  the  latter. 
G-ynandropsis  (Fig.  409)  is 
like  its  near  relative,  Cleome, 
except  that  this  very  long  stalk 
has  the  lower  part  of  the 
stamens  adnate  to  it :  the  in- 
ternode between  the  corolla 
and  calyx  is  broad  and  slightly 
elevated  (or  in  Cleome,  &c., 
narrower  and  longer)  ;  and 

408  409  8O  the  several  floral  circles 

are  as  it  were  spaced  apart  by  this  unusual  development  of 
receptacular  internodes.  In  Silene  (Fig.  408)  and  many  other 
plants  of  the  Pink  family,  an  internode  between  the  calyx  and 
corolla  is  prolonged  into  a  stalk  or  Stipe.1 

1  STIPE  is  the  general  name  of  a  stalk  formed  by  the  receptacle  or  some 
part  of  it,  or  by  a  carpel.  To  distinguish  its  particular  nature  in  any  case, 
the  following  terms  are  more  or  less  employed :  — 

THECAPHORE,  for  a  stipe  which  belongs  to  a  simple  pistil  itself  (where 
it  is  homologous  with  a  petiole),  and  is  no  part  of  the  receptacle,  as  in  Coptis 
or  Goldthread. 

GYNOPHORE,  where  the  stipe  is  an  internode  of  receptacle  next  below  the 
gynceciuin,  as  the  pod-stalk  in  some  Cruciferae,  Cleome,  and  Gynandropsis. 

GONOPHORE,  when  it  elevates  both  stamens  and  pistil,  as  it  seemingly 
does  in  the  lower  stipe  of  Gynandropsis,  Fig.  409. 

ANTHOPHORE,  when  the  stipe  is  a  developed  internode  between  the 
calyx  and  corolla,  as  in  the  Pink  family,  Fig.  408. 

GYNOBASE  is  a  term  properly  applied  to  a  short  and  comparatively  broad 
portion  of  receptacle  on  which  the  gynoecium  rests,  as  in  Rue  and  Orange 
(Fig.  414),  Houndstongue,  Sage,  &c.  This  may  extend  up  between  the  car- 
pels and  pass  into,  or  the  upper  part  become  a 

CARPOPHORE,  a  name  properly  applied  to  a  portion  of  receptacle  which 
is  prolonged  between  the  carpels  as  a  central  axis,  as  in  Geranium  (Fig. 
411)  and  many  Umbelliferae,  Fig.  412. 

FIG.  408.  Section  of  a  flower  of  Silene  Pennsylvania,  showing  the  stipe  or 
anthophore. 

FIG.  409.  Flower  of  Gynandropsis,  with  floral  circles  separated  on  the  elongated 
receptacle.^ 


FORMS   OF   THE   RECEPTACLE. 


213 


392.  Instead  of  forming  a  stalk,  the  elongation  may  be  continued 
between  the  carpels  in  the  form  of  a  slender  axis,  as  in  Gera- 
nium (Fig.  410,  411),  and  in  the  carpophore 
of  the  fruit  of  Umbelliferae,  Fig.  412.     In 
Geranium,    this    prolongation   of  receptacle 


extends  far  above  the  ovaries  as  a  beak,  to  which  the  styles  are 
adnate  for  most  of  their  length. 

393.  In  Nelumbium  (Fig.  413),  the  gynophore,  or  portion  of 
receptacle  above  the  stamens,  is  enlarged  into  a  singular  broadly 
top-shaped  body,  with  a  flat  summit,  in  which  the  pistils  (a  dozen 
or  more  isolated  carpels)  are  separately  immersed. 

394.  A  Disk  is  a  part  of  the  receptacle,  or  a  development  of  it, 
enlarged  under  or  around  the  pistil.     When  under  it  or  around 
its  base  and  free  from  the  calyx, 

the  disk  is  hypogynous,  as  in 
Orange,  Fig.  414.  Here  it  is  a 
kind  of  gynobase.  When  adher- 
ent to  or  lining  the  base  of  the 
calyx,  it  is  perigynous,  as  in 
Rhamnus  (Fig.  415,416)  and  Cherry  (Fig.  337)  :  when  carried 
by  complete  adnation  up  to  the  summit  of  the  ovary,  it  is  epigy- 
nous,  as  in  Cornus,  in  Umbelliferae,  &c.  Not  rarely  it  divides 
into  lobes,  as  in  Vitis  (Fig.  379,  380),  in  Periwinkle  and  most 
Apocynaceous  plants,  and  in  Cruciferae.  These  are  termed  glands 
of  the  disk,  and  indeed  are  commonly  glandular  or  nectariferous. 


FIG.  410.  Gyncecium  of  Geranium  maculatum.  411.  The  same  with  fruit  mature, 
the  five  ovaries  or  cells  and  the  lower  part  of  their  styles  separated  and  recurving  away 
from  the  prolongation  of  the  axis  or  receptacle,  to  which  they  were  at  flowering-time 
firmly  attached. 

FIG.  412.  Mature  fruit  of  Osmorrhiza,  the  two  carpels  splitting  away  below  from 
the  filiform  prolongation  of  the  receptacle,  or  carpophore. 

FIG.  413.  The  top-shaped  receptacle  of  Nelumbium,  with  the  pistils,  immersed  in 
hollows  of  its  upper  face. 

FIG.  415.  Flower  of  a  Rhamnus  or  Buckthorn,  and  416,  section  of  the  same,  show- 
ing a  thickened  perigynous  disk. 


214 


THE  FLOWER. 


It  is  not  possible  by  any  direct  demonstration  to  distinguish  be- 
tween such  productions  of  the  receptacle,  which  are  clawed  as 
belonging  to  the  axis,  and  suppressed  or  undeveloped  phyllous 
organs,  such  as  stamens,  which  glands  of  the  disk  may  some- 
times represent. 

395.  Hypanthium.  Inspection  of  Fig.  415,  416,  and  337,  and 
comparison  with  Fig.  339,  will  suggest  an  explanation  differ- 
ent from  that  which  is  generally 
adopted.  Instead  of  regarding  the 
calyx  as  beginning  on  a  level  with 
the  base  of  the  ovary,  and  the  cup 
as  lined,  more  or  less  thickly,  by 
an  expansion  of  the  receptacle  (the 
perigynous  disk) , 
the  catyx  may  be 
understood  to  begin 
where  this  and  the 
ovary  become  free 
from  each  other. 
Underthat  view,  the 
receptacle,  instead 
of  convex  or  protu 
berant,  is  here  con- 
cave, has  grown  up 
around  the  ovary, 
which,  however,  is  free  from  the  cup  in  the  earlier  cited  figures, 
but  immersed  in  it  in  Fig.  339  and  the  like.  A  comparison  with 
a  rose-hip,  an  apple,  and  a  pear  much  strengthens  this  interpre- 
tation, which  is  rather  largely  adopted  at  this  day,  at  least 
theoretically.  It  was  perhaps  first  proposed  by  Link,  who  intro- 
duced the  appropriate  name  of  HYPANTHUM.  A  hypanthium  or 
hypanthial  receptacle  is,  as  the  name  betokens,  a  flower-axis  or 
receptacle  developed  mainly  under  the  calyx.  The  name  is  a 
good  one,  in  any  case ;  and  such  structures  as  those  of  Catycan- 
thus  (Fig.  417-419),  a  rose,  a  pear  (the  lower  part  of  which  is 
evidently  an  enlargement  of  peduncle),  and  of  Cactus-flowers 
(Fig.  317),  although  quite  compatible  with  the  theory  of  adnation, 
are  more  simpty  explained  by  it.1 

1  But,  whether  the  cases  are  well  distinguishable  or  not,  it  by  no  means 
follows  that  the  receptacle  plays  such  a  part  in  all  instances  of  perigyny  and 
of  inferior  or  partly  inferior  ovary.  Such  a  view  is  attended  by  more  diffi- 
culties than  the  other.  Unless  the  mediation  of  an  invisible  receptacle  must 

FIG.  417.  Flowering  branch  of  Calycanthus.  418.  Vertical  section  of  the  urn-shaped 
receptacle,  the  imbricated  bracts  or  sepals  on  its  surface  cut  away.  419.  Mature 
fructiferous  receptacle  entire,  showing  some  scars  from  which  the  bracts  have  fallen. 


ADAPTATIONS   TO  FERTILIZATION.  215 

SECTION  IV.     CERTAIN  ADAPTATIONS  OF  THE  FLOWER  TO  THE 
ACT  OF  FERTILIZATION. 

§  1.     IN  GENERAL. 

396.  The  introduction  into  morphological  botany  of  the  con- 
siderations now  to  be  mentioned  should  have  dated  from  the 
year  1793,  in  which  Christian  Conrad  Sprengel  published  his 
curious  treatise  on  the  structure  of  flowers  in  special  reference 
to  insect  aid  in  their  fertilization.  For  this  book,  which  was 
wholly  neglected  and  overlooked  for  more  than  sixty  years,  con- 
tains along  with  some  fanciful  ideas  the  germs  of  the  present 
doctrine  and  many  excellent  illustrations  of  it.1  The  interest  in 
the  doctrine  now  prevalent  is  witnessed  by  a  copious  special 
literature,  beginning  with  the  publication,  in  1862,  of  Darwin's 
book  on  the  fertilization  of  Orchids  by  the  aid  of  insects.2 

be  invoked  whenever  there  is  a  junction  of  two  dissimilar  organs,  the  petals 
and  stamens  of  a  Lythrum  or  a  Cuphea  are  united  with  the  calyx  itself, 
instead  of  calyx  beginning  at  the  top  of  a  long  and  simple  tube.  And  if 
three  or  more  of  the  floral  whorls  may  be  congenitally  united,  why  not  these 
also  with  the  remaining  one  ?  Van  Tieghem,  in  his  Anatomie  Comparee  de 
la  Fleur,  maintains  wholly  the  old  view,  founding  it  upon  anatomical  struc- 
ture and  his  ability  to  trace  down  to  the  base  of  the  ovary  the  distinct 
vascular  bundles  of  the  several  involved  organs. 

1  C.  C.  Sprengel,  Das  entdeckte  Geheimniss  der  Natur  im  Bau  und  der 
Befruchtung  der  Blumen,  Berlin,  1793.     Even  earlier,  Koelreuter  ( Vorlaufige 
Nachricht,  etc.,  1761-1766)  recognized  the  necessity  of  insect-aid  to  various 
blossoms,  and  described  some  special  contrivances  for  the  purpose. 

2  Charles  Darwin,  On  the  Various  Contrivances  by  which  British  and 
Foreign  Orchids  are  fertilized  by  Insects,  and  on  the  Good  Effects  of  Inter- 
crossing,  London,  1862.    Ed.  2,  1877.  This  last  contains  a  list  of  the  papers 
and  books  which  bear  upon  the  subject,  published  since  1862. 

Other  leading  works  and  papers  on  the  subject  are,  exclusive  of  the 
other  volumes  and  papers  of  Darwin,  more  or  less  referred  to  hereafter. 

Treviranus,  Ueber  Dichogamie,  &c.,  in  Bot.  Zeitung,  xxi.  1863. 

Hugo  von  Mohl,  Einige  Beobachtungen  iiber  dimorphe  Bliithen,  Bot. 
Zeitung,  xxi.  1863. 

Delpino,  Pensieri  sulla  Biologia  Vegetale,  &c.,  1867.  Relazione  sull' 
Apparecchio  della  Fecondazione  nelle  Asclepiadie,  &c.,  1867.  Ulteriore 
Osservazioni  sulla  Dichogamia,  &c.,  1868-69,  1870,  and  later  papers. 

Axell,  Om  anordningarna  for  de  Fanerogama  Vaxternas  Befruchtung, 
Stockholm,  1869. 

Hildebrand,  Die  Geschlechter-Vertheilung  bei  den  Pflanzen,  1867,  and 
other  papers. 

Hermann  Miiller,  Die  Befruchtung  der  Blumen  dnrch  Insekten,  1873,  and 
papers  in  "  Nature  "  and  elsewhere. 

"  Flowers  and  their  Unbidden  Guests,"  an  English  translation  of  a  work 
by  Professor  Kerner,  which  describes  arrangements  in  blossoms  for  exclud- 
ing unwelcome  guests,  has  not  yet  reached  us.  It  introduces  the  new  terms 
Autogamy  and  Allogamy,  defined  on  the  following  page ;  the  latter  compre- 


216  THE  FLOWER. 

397.  The  subject,  here  considered  as  a  part  of  morphology, 
must  be  fully  treated,  as  regards  acts  and  processes,  under  physi- 
ology.    Every  thing  in  the  flower  is  in  relation  to  fertilization 
and  fructification,  directly  or  indirectly.     This  section  is  con- 
cerned with  those  adaptations  of  structure  by  means  of  which 
agents  external  to  the  blossom  are  brought  into  service  for  its 
fertilization. 

398.  Linnaeus  and  his  successors  taught  that  the  adjustments 
in  hermaphrodite  flowers  were  such,  on  the  whole,  as  to  secure  the 
application  of  the  pollen  of  its  stamens  to  the  stigma  of  its  pistil 
or  pistils.     The  present  view  is,  that  this  is  doubtless  strictly 
secured  in  certain  flowers  of  a  moderate  number  of  species,  but 
never  in  all  the  flowers  of  any  such  species ;  that  in  ordinary 
flowers,  where  it  may  commonly  take  place,  it  is  not  universal ; 
that  in  the  larger  number  of  species  there  is  something  or  other 
in  the  floral  structure  which   impedes  or  prevents   it.     Some 
flowers  are  adapted  for  close  fertilization ;  some  for  cross  fertili- 
zation ;  some  for  either.     Here  two  terms  need  definition,  viz. : 

Close  fertilization  or  Self-fertilization,  or  Autogamy,  the  appli ca- 
tion and  action  of  a  flower's  pollen  upon  its  own  pistil ; 

Gross  fertilization,  or  AUogamy,  the  action  of  the  pollen  of  one 
flower  on  the  pistil  of  some  other  flower  of  the  same  species. 
This  may  be  near,  as  when  between  flowers  borne  in  the  same 
cluster  or  on  the  same  plant ;  remote,  when  between  flowers  of 
distinct  plants  of  the  same  immediate  parentage  ;  most  remote, 
when  between  different  races  of  the  same  species.  Any  thing 
beyond  this  is  hybridization,  or  crossing  of  species. 

§  2.     ADAPTATIONS  FOR  ALLOGAMY  OR  INTERCROSSING. 

399.  The  doctrine  now  maintained  appears  to  have  been  first 
propounded  by  Sprengel  in  the  statement  that  "  Nature  seems 
to  have  wished  that  no  flower  should  be  fertilized  by  its  own 
pollen,"  —  a  proposition  which  is  not  wholly  tenable,  for  there 
are  blossoms  specially  adapted  to  self-fertilization.     It  was  re- 
affirmed in  our  day  by  Darwin,  in  a  similar  adage,  "Nature 
abhors  perpetual  self-fertilization,"  —  a  metaphorical  expression 
to  which  no  effective  exception  has  been  taken.     And  the  infer- 
ence was  drawn  by  him,  that  some  important  good  to  the  species 
must  result  from   propagation  through  the   union   of  distinct 
individuals,  and  especially  of  individuals  which  have  been  dis- 
tinct for  several  or  many  generations. 

bending  Geitonogamy,  fertilization  by  pollen  of  other  flowers  of  the  same 
plant,  and  Xenogamy,  by  pollen  from  a  flower  on  another  plant. 


ADAPTATIONS   FOR   INTERCROSSING.  217 

400.  The  actual  proposition,  simply  stated,  is  that  flowers  are 
habitually  intercrossed,  and  that  there  are  manifold  structural 
adaptations  which  secure  or  favor  intercrossing,  to  such  extent 
as  to  justify  the  proposition.     The  proof  of  the  proposition  is  an 
induction  from  a  very  great  number  of  particular  observations. 
That  intercrossing  is  beneficial  is  a  rational  inference  from  the 
array  of  special  adaptations  for  which  no  other  sufficient  reason 
appears,  or  (to  resume  the  metaphor)  from  the  vast  pains  which 
seem  to  have  been  taken  to  secure  this  end.     This  inference  has 
been  to  some  extent  confirmed  by  direct  experiment.1 

401.  Separation  of  the  sexes  is  a  direct  adaptation  to  inter- 
crossing, rendering  it  necessary  between  individuals  in  dioecious, 
and   largely  favoring  it  in  most  monoecious  and   polygamous 
flowers.     Strictly  close  fertilization  can  occur  in  hermaphrodite 
flowers  only  ;  but  it  is  in  these  that  the  most  curious  adaptations 
for  intercrossing  are  revealed. 

402.  The  agencies  to  the  one  or  the  other  of  which  most 
flowers  are  structurally  adapted  in  reference  to  intercrossing  are 
mainly  two  ;  viz.,  the  winds  and  animals,  of  these  chiefly  insects. 
Delpino  has  accordingly  classified  flowers  into  Anemophilous  and 
Entomophilous ;  literally  wind-lovers  and  insect-lovers,  but  de- 
noting  wind-fertilized   and   insect-fertilized,    according   to  the 
agent  by  which  pollen  is  transported.2    There  are  hermaphrodite 
and  unisexual  flowers  of  both  classes,  but  most  wind-fertilized 
flowers  are  unisexual. 

403.  Wind-fertilizable  or  anemophilous  flowers  are  mostly  neu- 
tral or  dull  in  color,  destitute  of  odor,  and  not  nectariferous. 
Their  principal  structural  adaptations  to  this  end,  besides  the 
separation  of  the  sexes  in  most  of  them,  are  the  superabundance, 
incoherency,  dryness,  and  lightness  of  the  pollen,  rendering  it 
very  transportable  by  wind  and  currents  of  air.     The  immense 
abundance  of  pollen,  its  lightness,  and  its  free  and  far  diffusion 
through  the  air  in  Pines,  Firs,  Taxodium,  and  other  Coniferae, 
are  familiar.     Their  pollen  fills  the  air  of  a  forest  during  anthe- 
sis ;   and  the  "showers  of  sulphur,"  popularly  so-called,  the 
yellow  powder  which  after  a  transient  shower  accumulates  as 
a  scum  on  the  surface  of  water  several  or  many  miles  from  the 

1  Darwin,  The  Effects  of  Cross  and  Self-Fertilization  in  the  Vegetable 
Kingdom,  London,  1876.     American  Edition,  New  York,  1877. 

2  Ornithophilous,  i.  e.  bird-fertilized,  flowers  are  to  be  ranked  with  entomo- 
philous.     The  large  blossoms  of  Trumpet  Creeper  (Tecoma  radicans)  and 
of  Trumpet  Honeysuckle  (Lonicera  sempervirens),  and  others,  are  commonly 
visited  and  probably  fertilized  by  humming-birds  as  well  as  by  moths  ;  and 
other  birds  are  known  to  play  a  similar  part  in  equatorial  regions. 


218  THE  FLOWER. 

nearest  source,  testifies  to  these  particulars.  All  amentaceous 
trees  (Willows  excepted) ,  Hemp,  Hops,  &c.,  are  wind-fertilized  ; 
and,  among  perfect  flowers,  those  of  most  Grasses,  Sedges,  and 
Plantago.  In  the  latter  families  especially,  the  anthers  are  pro- 
truded or  hung  out  in  the  air  only  when  just  ready  to  discharge 
their  pollen,  and  are  at  that  moment  suspended  on  suddenly 
lengthened  capillary  drooping  filaments,  fluttering  in  the  gentlest 
breeze ;  and  the  stigmas  are  either  dissected  into  plumes,  as  in 
most  Grasses,  or  beset  with  copious  hairs  on  which  pollen  is 
caught.  One  physiological  adaptation,  very  common  in  the  fol- 
lowing class,  is  not  unknown  among  hermaphrodite  wind-fertiliz- 
able  flowers,  where  it  is  important  for  securing  intercrossing,  viz. 
Dichogamy.  It  is  best  seen  in  the  common  species  of  Plantago 
or  Plantain,  and  is  described  below.  (408.) 

404.  Insect-fertilizable  or  entomophilous  flowers  are  correlated 
with  showy  coloration  (including  white,  which  is  most  showy  at 
dusk),  odor,  or  secretion  of  nectar,  often  by  all  three  modes  of 
attraction  to  insects  combined.     Some  insects,  moreover,  visit 
flowers  for  their  pollen,  a  highly  nutritious  article,  and  ordina- 
rily produced   in   such  abundance  that  much  may  be  spared. 
The  showiness  of  corolla  or  other  floral  envelopes  is  an  attractive 
adaptation  to  fertilization,  enabling  blossoms  to  be  discerned  at 
a  distance ;  nor  do  we  know  that  fragrance  or  other  scent  or 
that  nectar  subserves  any  other  uses  to  the  flower  than  that  of 
alluring  insects.     Adaptations  in  the  pollen  of  such  blossoms 
for  transportation  by  insects  are  various.     Commonly  the  grains 
are  slightly  moist  or  glutinous,   or  roughish,  or  studded  with 
projections,  or  strung  with  threads  (as  in  (Enothera) ,  so  as  not 
to   be   readily  dispersed   in   the  air,  but  to   have  some   slight 
coherence  as  well  as  capability  of  adhering  to  the  head,  limbs, 
or  bodies  of  insects,  especially  to  their  rough  surfaces;   and 
in  two  families  (Orchidaceae,  Asclepiadacese)  the  pollen  is  com- 
bined in  masses  and  with  special  adaptations  for  being  trans- 
ported  en    masse.    (421.)      With   this   the   stigma  is    usually 
correlated,  by  roughness,  moisture,  or  glutinosity.1 

405.  Adaptations  of  the  flower  itself  in  reference  to  insect 
visitation  are  wonderfully  various  ;  and  most  of  these  are  found 
upon  investigation  to  favor,  or  often  to  necessitate,  intercross- 
ing.    In  dioacious  flowers,  this  is  necessitated  by  the  separation ; 
in  monoecious  and  potygamous  flowers,  of  various  kinds  and 

1  Thus  nearly  every  Orchid  genus  but  one  has  a  persistently  glutinous 
stigma  ;  in  the  exceptional  one,  Cypripedium,  it  is  moist  and  minutely  rough- 
ened, in  correlation  with  the  loosely  granular  or  pultaceous  pollen  which  it 
is  to  receive. 


ADAPTATIONS   FOR  INTERCROSSING.  219 

rjagrees  of  separation,  pollen  is  very  commonly  borne  from 
plant  to  plant ;  in  hermaphrodite  flowers  only  are  more  special 
arrangements  needed  to  secure  intercrossing  or  a  certain  measure 
of  it,  and  in  these  such  arrangements  abound. 

406.  Irregularity  is  one  of  the  commonest  modifications  of  the 
flower  (326,  337)  :  it  is  never  conspicuous  except  in  blossoms 
visited  by  insects  and  generally  fertilized  by  their  aid ;  and  it 
finds  rational  explanation  on  the  score  of  utility  in  this  regard.1 

407.  Dichogamy,  a  term  introduced  by  C.  C.  Sprengel,  who 
first  noticed  and  described  it,  is  one  of  the  most  usual  and  effect- 
ual (rather  physiological  than  morphological)  adaptations  for  the 
promotion  of  intercrossing  between  hermaphrodite  flowers.     It 
means  that  such  intercrossing  is  brought  to  pass  by  a  difference 
in  the  time  of  maturity  of  anthers  and  stigma ;  this  rendering 
dichogamous  blossoms  practically  the  same  as  dioecious  or  mon- 
oecious in  respect  to  fertilization,  while  there  is  the  economical 
gain  that  all  the  flowers  are  fertile.     According  to  whether  the 
anthers  or  the  stigmas  are  precocious,  dichogamous  flowers  are 

Proterandrous  (or  Protandrous) ,  when  the  anthers  mature  and 
discharge  their  pollen  before  the  stigma  of  that  blossom  is  recep- 
tive of  pollen ; 

Proterogynous  (or  Protogynous) ,  when  the  stigmas  are  in 
receptive  condition  before  the  anthers  have  matured  their  pollen. 

Synanthesis*  the  maturing  of  the  two  sexes  simultaneously  or 
nearly  so,  is  however  made  to  secure  the  same  result  through 
special  arrangements. 

408.  Proterogyny.     The  Plantains,  such  as  Plantago  major  and 
P.  lanceolata,  are  familiar  instances  of  this  in  a  wind-fertilized 
genus  with  hermaphrodite  flowers.     The  anthesis  proceeds  from 
base  to  apex  of  the  spike  in  regular  order,  and  rather  slowly. 
While  the  anthers  are  still  in  the  unopened  corolla  and  on  short 
filaments,  the  long  and  slender  hairy  stigma  projects  from  the  tip 
and  is  receiving  pollen  blown  to  it  from  neighboring  plants  or 

1  This  did  not  escape  the  attention  of  Sprengel  in  the  last  century,  and 
along  with  it  the  fact  that  strictly  terminal  and  also  vertical  flowers,  whether 
erect  or  suspended,  are  seldom  irregular,  while  comparatively  horizontal  or 
obliquely  set  flowers  more  commonly  are  so.     The  irregularity  is  in  refer- 
ence to  a  landing  place  for  the  visiting  insect,  or  also  to  storage  of  or  accessi- 
bility to  nectar,  &c. 

Darwin  (Forms  of  Flowers,  147)  remarks  that  he  does  not  know  of  a 
single  instance  of  an  irregular  flower  which  is  wind-fertilized. 

2  Synacmy  is  the  term  proposed  by  A.  W.  Bennett,  in  Journal  of  Botany, 
viii.  (1870),  316,  with  its  opposite,  Heteracmy,  f or  proterandry  and  proterogyny. 
The  latter  names,  in  their  shorter  form  (protandry  and  protogyny),  appear  to 
have  originated  with  Hildebrand.  1867. 


220  THE   FLOWEB. 

spikes  :  a  day  or  two  afterwards,  the  corolla  opens,  the  filaments 
greatly  lengthen,  and  the  four  anthers  now  pendent  from  them 
give  their  light  pollen  to  the  wind  ;  but  the  stigmas  of  that  flower 
and  of  all  below  it  on  that  spike  are  withered  or  past  receiving 
pollen.  Among  Grasses,  Anthoxanthum  is  in  the  same  case. 
The  arrangement  is  somewhat  similar  to  the  Plantain  in  Amor- 
pha,  which  is  fertilized  by  insects,  the  simple  stigma  projecting 
beyond  the  corolla  in  bud,  while  the  anthers  are  still  immature 


and  enclosed.  Scrophularia  is  a  good  instance  of  proterogyny 
in  flowers  fertilized  by  bees.  The  flower  is  irregular  (Fig. 
420—422) ,  and  is  approached  from  the  front,  the  spreading  lower 
lobe  being  the  landing  place.  Fig.  420  represents  a  freshly 
opened  blossom ;  and  Fig.  421,  a  section  of  it.  Only  the  style 
tipped  with  the  stigma  is  in  view,  leaning  over  the  landing  place  ; 
the  still  closed  anthers  are  ensconced  below.  The  next  day  or 
a  little  later  all  is  as  in  Fig.  422.  The  style,  now  flabby,  has 
fallen  upon  the  front  lobe,  its  stigma  dry  and  no  longer  receptive : 
the  now-opening  anthers  are  brought  upward  and  forward  to  the 
position  which  the  stigma  occupied  before.  A  honey-bee,  taking 
nectar  from  the  bottom  of  the  corolla,  will  be  dusted  with  pollen 
from  the  later  flower,  and  on  passing  to  one  in  the  earlier  state 
will  deposit  some  of  it  on  its  fresh  stigma.  Self-fertilization 
here  can  hardly  ever  take  place,  and  only  through  some  disturb- 
ance of  the  natural  course. 

409.  Proterandry.  The  process  is  the  reverse,  and  is  at- 
tended with  much  more  extended  movements  in  Clerodendron 
Thompson!  ae,  a  Verbenaceous  tropical  African  climber  now  com- 
mon in  conservatories.  The  adaptations  in  this  flower  (which 
we  indicated  long  ago)  are  exquisite.  The  crimson  corolla 
and  bright  white  calyx  in  combination  are  very  conspicuous. 
The  long  filiform  filaments  and  style,  upwardly  enrolled  in  the 

FIG  420,  421.  Early  opened  flower  of  Scrophularia  nodtsa,  and  a  longitudinal 
section.  422.  Flower  a  day  or  two  later. 


ADAPTATIONS  FOE,   INTERCROSSING.  221 

bud,  straighten  and  project  when  the  corolla  opens  :  the  stamens 
remain  straight,  but  the  style  proceeds  to  curve  downward  and 
backward,  as  in  Fig.  423.  The  anthers  are  now  discharging 
pollen  :  the  stigmas  are  immature  and  closed.  Fig.  424  repre- 
sents the  flower  on  the  second  day,  the  anthers  effete,  and  the 
filaments  recurved  and  rolled  up  spirally ;  while  the  style  has 


taken  the  position  of  the  filaments,  and  the  two  stigmas  now 
separated  and  receptive  are  in  the  very  position  of  the  anthers  the 
previous  day.  The  entrance  by  which  the  proboscis  of  a  butterfly 
ma}*  reach  the  nectar  at  bottom  is  at  the  upper  side  of  the  orifice. 
The  flower  cannot  self-fertilize.  A  good-sized  insect  flying  from 
blossom  to  blossom,  and  plant  to  plant,  must  transport  pollen 
from  the  one  to  the  stigma  of  the  other. 

410.  Proterandry  abounds  among  common  flowers.  It  is 
conspicuous  in  Gentians  and  in  nearly  all  that  family.  But, 
while  in  Gentians  the  short  style  is  immovable  and  erect,  in 
Sabbatia  it  is  thrown  strongly  to  one  side,  out  of  the  way  of  and 
far  below  the  stamens,  the  branches  closed  and  often  twisted,  so 
that  the  stigma  is  quite  inaccessible  until  the  stamens  have  shed 
their  pollen  :  then  the  style  becomes  erect,  untwists,  its  two  flat 
branches  separate,  and  expose  the  stigmatic  surface  of  their  inner 
face  in  the  place  which  the  anthers  occupied.  In  Sabbatia 
angularis,  Lester  F.  Ward*  observed  that  the  anthers  of  freshly 

*  In  Meehan's  Gardeners'  Monthly,  September,  1878,  278. 
FIG.  423.    Flower  of  Clerodendron  Thorn psoniae,  first  day;  424,  second  day. 


222  THE   FLOWER. 

opened  blossoms  are  all  thrown  to  one  side  almost  as  strongly 
as  the  style  is  thrown  in  the  opposite  direction.  One  of  our 
common  Fireweeds,  Epilobium  angustifolium  or  E.  spicatum,  as 
it  is  variously  called,  which  is  common  all  round  the  northern 
hemisphere,  is  similar  to  Sabbatia  in  behavior.  In  the  freshly 
opened  flower,  while  the  anthers  are  in  good  condition  and  are 


giving  their  pollen  to  bees,  the  still  immature  style  is  strongly 
curved  downward  and  backward,  as  in  Fig.  425.  Two  or  three 
days  later,  when  the  pollen  is  mostly  shed,  the  style  straightens, 
lengthens  to  its  full  dimensions,  and  spreads  its  four  stigmas  over 
the  line  of  the  axis  of  the  blossom  (Fig.  426),  in  the  very 
position  to  be  pollinated  by  a  bee  coming  from  an  earlier  flower. 
411.  In  the  following  instances  of  proterandry,  the  style  is 
made  the  instrument  of  distributing  the  pollen  which  it  is  not 

itself  to  use.  The 
anthers  of  a  Cam- 
panula discharge  all 
their  pollen  in  the 
unopened  bud,  and  it 
is  nearly  all  deposited 
on  the  style  which 
they  surround,  the 
upper  part  of  which 
is  clothed  with  a  coat 
of  hairs  for  holding 

„  „  the  pollen.  (Fig.  427.) 

In  the  open  flower,  the  stamens  are  found  to  be  empty  and  withered, 
as  in  Fig.  428.  These  flowers  are  visited  by  bees  and  other  insects 
for  the  pollen.  While  this  is  going  on,  and  while  the  pollen  is 
fresh  and  plentiful,  no  stigma  is  apparent.  Later,  the  top  of  the 
style  opens  into  three  (in  some  species  five)  short  and  spreading 
branches,  the  inner  faces  of  which  are  the  stigmas.  Although 

FIG.  425,  426.  Flowers  of  Epilobium  angustifolium  or  spicatum ;  in  the  first, 
freshly  expanded ;  in  the  second,  a  few  days  older. 

FIG.  427.  Vertical  section  of  an  unopened  flower  of  Campanula  rapnnculoldes : 
the  broad  white  lines  are  sections  of  two  anthers.  428.  Same  of  an  older  flower. 


ADAPTATIONS  FOR  INTERCROSSING. 


223 


so  close  at  hand,  little  if  any  of  the  pollen  of  that  flower  can 

reach  the   stigmas.      These   actually  get   fertilized  by  pollen 

brought  by  bees,  which  come  loaded  with  it  from 

other  flowers  and  other  plants.    Symphyandra  differs 

from   a   true   Campanula   chiefly  in  the   continued 

cohesion  of  the  five  anthers  into  a  tube  around  the 

style.  (Fig.  429,  430.)    The  pollen  is  discharged  on 

and  held  by  the  hairy  upper  portion  of  the  style. 

Soon  after,  the  corolla  expands,  the  lower 

part  of  the  st}7le  lengthens,  and  carries 

the  pollen-loaded  part  out  of  and  above 

the  anther-tube,  as  in  Fig.  430  ;  lastly,  the 

three  connivent  tips  of  the  style  diverge 

and  expose  the  stigmas  to  pollen  mainly 

brought  by  bees  from  other  flowers.    By  a 

slight  further  modification  in  Lobelia  and 

in  Compositae,  pollen  is  pushed  out  of  the 

anther-tube  by  the  tip  of  the  style  as  it 

lengthens,  or  by  the  very  back  of  the  two 

stigmas,  the  faces  of  which,  afterwards 

exposed,  are  not  to  receive  this,  but  other 

pollen,  though  it  may  at  times  receive  some  of  its  own.     The 

arrangement  in  Compositae  is  here  illustrated  from  Leptosyne 

maritima  (Fig.  431-435),  a  showy  plant  of  Southern  California, 

now  not  very  rare  in  cultivation.     The  large  flowers  around  the 


FIG.  429.    Stamens  and  pistil  of  a  young,  and  430,  same  from  an  old  flower  at 
Symphyandra  pendula 

FIG.  431.    Head  of  flowers  of  Leptosyne  maritima,  of  the  natural  size. 


THE  FLOWER. 


margin  (ray-flowers,  with  ligulate  corolla),  one  of  which  is  sepa- 
rately shown  in  Fig.  432,  are  pistillate  only :  the  enlarged  and 

extended  open  part  of  the 
corolla  (bright  yellow  in 
color)  serves  for  attrac- 
tion, the  circle  of  ra,ys 
gives  the  appearance  as 
of  a  single  large  flower. 
The  flowers  of  the  disk 
or  whole  central  part  are 
hermaphrodite,  and  with 
narrow  tubular  corollas, 
from  the  orifice  of  which 
•  projects  the  greater  part 
of  the  tube  of  five  co- 

alescent  anthers.  The  pollen  is  early  discharged  into  the  interior 
of  this  tube.  The  style,  with  somewhat  enlarged  and  brush-like 
tip,  at  first  reaches  only  to  the 
bottom  of  the  anther-tube:  it 
slowly  lengthens,  pushes  the 
pollen  before  it  out  of  the  tube 
(Fig.  433)  and  into  the  way  of 
insects  of  various  kind,  which, 
travelling  over  the  surface,  con- 
vey it  to  older  flowers  of  the  same 


head  and  of  other  plants.  The  style,  elongating  yet  more,  raises 
some  of  the  pollen  still  higher  (as  in  Fig.  434)  ;  and  at  length  its 
two  branches  separate  and  diverge  (Fig.  435) ,  exposing  to  other 
pollen  the  stigmatic  receptive  surface  which  until  now  was  un- 
approachable. 

412.  In  Parnassia,  which  has  sessile  stigmas,  their  receptive 
surface  is  actually  not  formed  until  the  anthers  become  effete ; 

FIG.  432.  A  ligulate  female  flower  of  the  same,  and  a  central  hermaphrodite  flower 
433.  Upper  part  of  the  latter,  more  enlarged,  the  tube  of  anthers  projecting  from  the 
corolla,  and  the  pollen  projecting  from  apex  of  the  anther-tube,  being  pushed  up  by  the 
lengthening  of  the  style  beneath.  434.  This  style  now  projecting,  and  some  pollen  still 
resting  on  its  tip.  435.  Tip  of  same  style  (more  advanced  and  magnified);  the  two 
branches  spreading,  still  carrying  some  pollen  on  the  apex  of  each  arm  or  branch,  and 
by  the  divergence  now  exposing  the  stigmatic  inner  faces.. 


ADAPTATIONS   FOB   INTERCROSSING. 


225 


and,  as  the  plants  or  stems  are  single-flowered,  they  are  function- 
ally dioecious  while  structurally  hermaphrodite. 

413.  The   adaptations   for  hermaphrodite  intercrossing  with 
synanthesis  (407) ,  »'.  e.  where  there  is  no  essential  difference  of 
time  in   the   maturing  of  anthers   and   stigma,    are   manifold. 
They  may  be  classed  into  those  without  and  those  with  dimor- 
phism of  stamens  and  pistils,  or,  in  other  words,  those  with 
Homogonous  and  those  with  Heterogonous  flowers.1 

414.  The   cases  without  dimorphism   are   the  most  various, 
certain  families  having  special  types ;  and  are  of  all  degrees, 
from  those  that  require  intercrossing  to  those  that  merely  favor 
or  permit  it.     For  the  present  purpose,  having  only  morphology 
in  view,  it  suffices  to  bring  to  view  two  or  three  cases  or  types  of 

415.  Particular  Adaptations  in  hermaphrodite  blossoms,  not 
involving  either  dichogamy  or  dimorphism.     These  are  exceed- 
ingly various  ;  but  they  may  be  distinguished  into  two  general 
kinds,  namely:    1,  where  loose  and 

powdery  pollen  is  transported  from 
blossom  to  blossom  in  separate  grains, 
and  2,  where  pollen-masses  or  the 
whole  contents  of  anthers  are  bodily 
so  transported. 

416.  Papilionaceous    flowers   (such 
as  pea-blossoms,  338) — having   ten 
stamens  enclosed  with  a  single  pis- 
til in  the  keel, 

of  the  corolla, 

their  anthers  in 

close  proximity 

to  the  stigma  — 

were    naturally 

supposed  to  be 

self-fertilizing ;   and  so  they  sometimes  are,  yet  with  marked 

adaptations  for  intercrossing.     None  are  less  so  than  those  of 

1  Terms  proposed  in  Amer.  Jour.  Sci.  ser.  3,  xiii.  82,  and  in  Amer. 
Naturalist,  January,  1877.  Dimorphism  in  flowers  may  affect  the  perianth 
only,  and  not  the  yov-fi  or  essential  organs ;  or  there  may  be  two  kinds  of 
flowers  as  respects  these  also,  but  with  no  reciprocal  relations,  as  in  cleisto- 
gamous  dimorphism  (534) ;  or  of  two  kinds  essentially  alike  except  in  stamens 
and  pistil,  and  these  reciprocally  adapted  to  each  other,  which  is  heterogonous 
dimorphism,  or,  when  of  three  kinds,  trimorphism. 

FIG.  436.  Flower  of  Wistaria  Sinensis,  natural  size.  437.  Same  enlarged,  with 
standard,  wings,  and  half  the  keel  removed.  438.  Same  with  the  keel  depressed,  as  it 
is  when  a  bee  alights  on  this  its  usual  landing  place,  the  cluster  of  anthers  and  stigma 
thus  brought  up  against  the  bee's  abdomen.  439.  Style  and  stigma,  with  part  of  the 
ovary,  more  magnified,  a  fringe  of  fine  bristles  around  the  stigma. 
15 


THE  FLOWEK. 


Wistaria  (Fig.  436-439),  in  which  the  light  fringe  of  stiff  hairs 
around  the  stigma  (shown  in  Fig.  439)  would  not  prevent  pollen 
of  surrounding  anthers  from  falling  upon  it.  Yet  when  a  bee 

alights  upon  the  keel, 
with  head  toward  the 
base  of  the  flower,  and 
proboscis  is  inserted 
for  nectar  between  the 
foot  of  the  standard 
and  the  keel,  the  latter 
is  depressed  by  the 
weight,  so  that  the  ab- 
domen of  the  insect  is 
brought  against  the  ten 
anthers  and  the  stig- 
ma, becoming  thereby 
smeared  with  pollen, 
some  of  which  when 
other  blossoms  are  vis- 
ited cannot  fail  to  be  applied  to  their  stigmas.  The  very  similar 
flower  of  Locust  (Robinia) ,  like  that  of  the  Pea,  adds  an  adapta- 
tion in  favor  of  intercrossing.  The  style  for  some  length  below  the 
stigma  is  covered  with  a  short  beard  of  hairs,  as  is 
seen  in  Fig.  442.  The  anthers  open  early  and  dis- 
charge their  pollen,  which  mainly  lodges  on  this 
beard  (Fig.  443) ,  in  a  manner  which  may  thus  far 
be  likened  to  the  case  of  Campanula.  (411.)  The 
wings  and  the  keel  are  yoked  together,  and  are 
together  depressed  by  the  weight  of  an  alighting 
443  bee.  This  does  not  bring  out  the  anthers  as  in 

Wistaria,  but  these  remain  until  effete  within  the  sac,  while  the 
stigma  and  the  pollen-laden  part  of  the  style  (Fig.  441)  are 
projected  against  the  bee's  abdomen,  which,  by  the  oblique 
movement,  is  first  touched  by  the  stigma  and  next  brushed  over 
with  pollen  by  the  style  below.  So  that,  in  visiting  a  succession 
of  blossoms,  some  pollen  of  one  flower  is  transferred  to  the  body 
of  the  bee,  and  thence  to  the  stigma  of  the  next  flower,  which 
flower  immediately  gives  to  the  same  spot  some  of  its  pollen,  to 
be  transferred  to  the  next  flower's  stigma,  and  so  on. 

417.  Two   special  modifications  of  the   papilionaceous  type 

FIG.  440.  Flower  of  Robinia  hispida,  the  standard  and  wings  removed.  441.  Same, 
as  depressed  by  the  weight  of  a  bee,  causing  the  stigma  and  pollen-laden  tip  of  the  style 
to  protrude.  442.  Enlarged  section  of  same  in  the  bud,  leaving  one  keel-petal,  half  the 
stamens,  and  the  pistil  in  view.  443.  Style  and  stigma  at  a  later  period,  the  beard 
loaded  with  pollen ;  more  magnified. 


ADAPTATIONS    FOB,   INTERCROSSING. 


227 


need  particular  mention.  One  of  them,  the  Bean-blossom,  is 
well  known  to  botanists ;  the  other  not  so.  The  peculiarity  in 
the  common  Bean,  Fhaseolus  vulgaris,  and  its  nearest  relatives, 
is  that  the  keel,  enclosing  the  stamens  and  pistil,  is  prolonged 


into  a  narrow  snout  which  is  spirally  coiled  (as  in  Fig.  444-446)  ; 
that  the  stigma  is  oblique  on  the  tip  of  the  style,  and  the  beard 
on  the  style  is  mainly  on  the  same  side  that  the  stigma  is  :  the 
wing-petals  stand  forward  and  turn  downward,  forming  a  con- 
venient landing  place  for  bees.  As  in  the  Locust-blossom,  the 
anthers  early  discharge  their  pollen,  much  of  which  adheres 


lightly  to  the  beard  of  the  style.  In  the  untouched  flower,  all 
from  first  to  last  is  concealed  in  the  coiled  keel.  Press  down  the 
wing-petals,  and  first  the  stigma  and  then  the  pollen-laden  tip 
of  the  style  projects  from  the  orifice  :  remove  the  pressure,  and 
they  withdraw  within.  When  this  pressure  is  made  by  a  bee, 
resting  on  the  wing-petals  while  searching  for  nectar  within  the 
base  of  the  blossom  between  the  keel  and  the  standard,  the  same 
movement  occurs :  the  stigma  first,  and  then  the  pollen  on  the 
style,  strikes  against  a  certain  portion  of  the  front  or  side  of 
the  bee's  body,  and  the  repetition  of  this  operation  causes  the 
fertilization  of  each  blossom  by  other  than  its  own  pollen.  A 
slighter  pressure  or  lighter  movement  of  the  wing-petals  suffices 

FIG.  444.  Flower  of  Garden  Bean,  Phaseolus  vulgaris.  445  Same  with  wing- 
petals  pressed  down  and  tip  of  style  projecting  from  the  orifice  of  the  keel.  446.  Same 
as  444  enlarged,  and  standard  and  wings  removed.  447.  Upper  part  of  keel,  in  the 
condition  of  445,  enlarged,  showing  plainly  the  projecting  style.  448  Section  of  the 
keel,  enlarged,  showing  the  style  within  before  the  anthers  open :  stamens  for  sake  of 
clearness  not  delineated.  449.  Pistil  detached  from  an  older  flower;  the  brush  loaded 
with  pollen. 


THE   FLOWER. 


to  jostle  some  of  the  pollen  down  upon  its  own  stigma,  so  that 

self-fertilization  is  not  uncommon. 

418.  Apios  tuberosa,  a  near  relative  of  Phaseolus,  exhibits  a 

different  and  equally  curious  modification  of  the  same  parts. 
The  wing-petals  for  landing  place 
are  similar :  the  standard  is  pro- 
portionally large,  firm  in  texture, 
and  shell-shaped  or  concave,  with 
a  small  boss  at  the  tip  as  seen  from 
behind,  or  a  shallow  sac  as  seen 
450  ^  from  the  front :  the  keel  is  narrow 

and  sickle-shaped  ;  it  arches  across  the  front  of  the  flower,  and 

the  blunt  apex  rests  in  the  notch  or  shallow  sac  of  the  tip  of  the 


standard.  (Fig.  450, 452, 453.)  So  it  remains  if  untouched  until 
the  blossom  withers :  no  self-fertilization  has  ever  been  observed, 
and  none  ordinarily  occurs.  The  anthers 
are  assembled  close  around  the  stigma, 
but  a  little  short  of  it  (Fig.  452)  ;  the 
pollen  is  not  early  nor  copiously  shed  in 
the  enclosure :  the  small  terminal  stigma 
is  at  first  covered  with  a  pulpy  secretion, 
which  at  length  collects  into  a  soft  ring 
around  its  base  over  or  through  which  no 
pollen  passes.  But  when  the  keel  is  liber- 
ated by  lifting  from  underneath,  it  curves 
promptly  into  the  shape  shown  in  Fig. 

FIG.  450.  Flower  of  Apios  tuberosa,  unvisited.  451.  Same  after  visitation,  the 
keel  dislodged  from  the  retaining  notch,  and  more  incurved;  the  tip  of  the  style  pro- 
truded and  thrust  forward,  followed  by  the  anthers. 

FIG.  452.  Enlarged  vertical  section  of  flower-bud  of  Apios  tuberosa.  453.  A  flower 
with  half  the  standard  cut  away,  to  show  the  blunt  apex  of  the  keel  resting  in  the 
notcfc.  454.  Diagram  of  flower,  with  half  of  the  standard  cut  away,  to  show  what  takes 
place  when  the  apex  of  the  keel  is  liberated.  The  figures  (also  those  from  423  to  the 
present),  and  the  first  account  of  the  adaptations  of  Apios,  were  published  in  the  Amer- 
ioan  Agriculturist  in  1876. 


ADAPTATIONS   FOR   INTERCROSSING.  229 

451,  or  better  in  Fig.  454,  where  the  dotted  lines  indicate  its 
original  position ;  and  first  the  end  of  the  style,  tipped  with 
its  stigma,  is  pushed  forward,  and  then  the  anthers  come  into 
view.  The  flowers  are  visited  by  humble-bees,  and  sometimes 
by  honey-bees.  In  searching  for  nectar  at  the  base  of  the  flower, 
they  probably  push  forward  into  the  space  under  the  arching 
keel,  and  by  slightly  elevating  dislodge  its  apex  ;  when  first  the 
stigma  and  then  the  anthers  are  brought  against  some  portion 
of  the  insect's  body,  and  against  the  same  portion  in  succeeding 
blossoms,  thus  effecting  cross-fertilization.  This  rationally  ex- 
plains a  remarkable  adaptation,  which  seems  to  be  not  otherwise 
intelligible. 

419.  Special   Adaptations.      Two  of  these,  each  peculiar  to 
the  genus,  may  here  be  referred  to.     In  Kahnia-blossoms  (Fig. 


455-458),  the  anthers  discharge  the  pollen  through  a  small 
orifice  at  the  apex  of  each  cell,  in  this  respect  agreeing  with 
Rhododendrons  and  their  other  relatives ;  but 
none  of  them  utilize  this  family  peculiarity  in  the 
manner  of  Kalmia.  In  the  flower-bud,  each  of  the 
ten  anthers  is  lodged  in  a  small  cavity  or  pocket 
(externally  a  boss)  of  the  corolla,  in  a  way  analo- 
gous to  that  in  which  the  keel  of  Apios  is  lodged  in 
the  tip  of  the  standard  (418)  :  the  expansion  of  the 
border  of  the  corolla  in  anthesis  curves  the  fila- 
ments outward  and  backward  ;  and  when  the  bowed  453 
stamens  are  liberated  by  rough  jostling  they  fly  up  elastically, 
and  the  pollen  is  projected  from  the  two  orifices.  Some  pollen 
may  possibly  be  thrown  upon  the  single  small  stigma  at  the 
tip  of  the  style,  which  rises  much  above  the  stamens.  But  the 
anthers  are  not  dislodged  when  undisturbed,  at  least  until 
after  the  elasticity  of  the  filaments  is  lost :  they  are  dislodged  by 
humble-bees,  which  circle  on  the  wing  over  the  blossom,  the 

FIG.  455  Vertical  section  of  a  flower-bud  of  Kalmia  latifolia,  showing  the  anthers 
lodged  in  the  pockets  of  the  corolla.  456.  Expanded  flower,  with  bowed  stamens. 
457.  Vertical  section  of  the  same.  458.  A  stamen,  enlarged. 


230 


THE  FLOWER. 


under  side  of  the  abdomen  frequently  touching  the  stigma,  while 
the  proboscis  is  searching  round  the  bottom  of  the  flower,  liberat- 
ing the  stamens  in  the  process,  which  one  by  one  project  their 
pollen  upon  the  under  side  of  the  insect's  body.  In  the  passage 
from  flower  to  flower,  pollen  is  thus  conveyed  from  the  anthers 
of  one  to  the  stigma  of  another. 

420.  Iris  has  three  stamens,  one  before  each  sepal  or  outer 
lobe  of  the  perianth,  and  behind  each  petal-like  lobe  of  the  style 

(Fig.  459)  :  the  stigma,  a 
shelf-like  plate  of  each  lobe, 
is  just  above  the  anther ; 
but,  as  the  anther  faces 
outward  and  the  stigma  is 
higher  and  faces  inward,  no 
pollen  can  find  its  way  from 
the  one  to  the  other.  But  the 
adaptation  of  parts  is  admir- 
able for  conveyance  by  bees, 
which,  standing  upon  the 
only  landing  place,  the  re- 
curved sepal,  thrust  the  head 
down  below  the  anther,  and 
in  raising  it  carry  off  pollen, 
to  be  afterwards  lodged 
upon  the  stigmas  of  other 
flowers  which  they  visit. 

421.  Transportation  of  Pollinia,   or  of  all  the  pollen  in  a 
mass,  is  effected  in  most  of  the  species  of  two  large  orders, 
not  otherwise  allied,  the  Asclepiadacese  and  the  Orchidacese. 
While  in  the  Iris  family  the  number  of  stamens  is  reduced  from 
six  to  three,  in  all  the  Orchis  family,  except  Cypripedium,  the 
stamens  are  further  reduced  to  a  single  one ;  but  the  pollen  is 
peculiarly  economized.     That  of  Arethusa  is  in  four  loose  and 
soft  pellets,  in  an  inverted  casque-shaped  case,  hinged  at  the 
back,  resting  on  a  shelf,   the  lower  face  of  which  is  glutinous 
stigma,  over  the  front  edge  of  which  the  casque-shaped  anther 
slightly  projects  ;  and  this  anther  is  raised  by  the  head  of  a  bee 
when  escaping  out  of  the  gorge  of  the  flower.    The  loose  pellets 
of  pollen  are  caught  upon  the  bee's  head,  to  the  rough  sur- 
face of  which  they  are  liable  to  adhere  lightly  and  so  to  be  carried 
to  the  flower  of  another  individual,  there  left  upon  its  glutinous 

FIG.  469.  Flower  of  Iris  pumila,  with  front  portion  and  half  of  one  petaloid  style- 
lobe  and  stigma  cut  away.  The  section  of  the  stigma  is  seen  edgewise:  the  rough 
upper  surface  only  is  stigmatic. 


ADAPTATIONS   FOR  INTERCROSSING. 


231 


stigma  by  the  same  upward  movement  which  immediately  after- 
ward raises  the  anther-lid  and  carries  away  its  pollen,  to  be 
transferred  to  a  third  blossom,  and  so  on. 

422.  But  it  is  in  Orchis 
and  in  the  commoner  re- 
presentatives of  Orchis  in 
North  America  (viz.  Ha- 
benaria,  &c.)  that  the 
most  exquisite  adapta- 
tions are  found,  and  the 
greatest  economy  se- 
cured ;  paralleled,  how- 
ever, by  most  of  the  very- 
numerous  and  various  epi- 
phytic and  by  various  ter- 
restrial Orchids  of  warmer 
regions.  A  single  illus- 
tration may  here  suffice ; 
and  Darwin's  volume  on 
the  Fertilization  of  Or- 
chids (396,  note),  with 
its  references  to  the 
copious  literature  of  the 
subject,  may  be  studied 
for  full  particulars  and 
their  bearings.  The  flower 
is  trimerous,  and  the  peri- 
anth adnate  to  the  ovary, 
therefore  apparently  de- 
veloped upon  its  sum- 
mit. The  three  external 
parts  of  the  perianth, 
which  in  Habenaria  orbi- 
culata  (Fig.  460)  are 
much  the  broader,  are  the 
sepals:  the  three  alternate 
and  internal,  the  petals  : 
the  base  of  the  long  and  narrow  petal  which  is  turned  downward 
is  hollowed  out  and  extended  below  into  a  long  tube,  closed  at 
bottom,  open  at  top  (the  spur  or  nectary),  in  which  nectar  is 


FIG.  460.  Flower  of  Habenaria  or  Platanthera  orbiculata,  enlarged.  461.  Combined 
stamen  and  stigma,  more  enlarged.  462.  One  of  the  two  pollen-masses  (pollinia),  with 
its  stalk  and  glutinous  disk  or  glaud.  462«.  Lower  part  of  this  stalk  and  ita  diak,  more 
magnified. 


282 


THE  FLOWEK. 


copiously  secreted  and  contained.  The  central  part  of  the 
blossom,  beyond  the  orifice  of  the  nectary  (shown  separately  in 
Fig.  461),  consists  of  one  anther  and  a  stigma,  fused  together 
(the  clinandrium)  :  the  marginal  portions,  opening  by  a  long 
chink,  are  the  two  cells  of  the  anther,  approximate  at  their 
broader  portion  above,  widely  divergent  below :  most  of  the 
lower  part  of  the  space  between  is  excessively  glutinous,  and  is 
the  stigma.  The  grains  of  pollen  are  united  by  means  of  short 
threads  of  very  elastic  tissue  into  small  masses,  and  these  into 
larger,  and  at  length  into  pellets,  having  stalks  of  the  same 
elastic  tissue,  by  which  they  are 
all  attached  to  a  firmer  central 
stalk,  or  caudicle.  (Fig.  463-465.) 
To  the  lower  end  of  this  caudicle 
(directly  to  the  end  of  it  in  our 
Habenariae  and  Orchises  gener- 
ally, in  this  instance  to  the  inner 
side  of  the  end,  with  a  thick  inter- 
mediate base  intervening),  is  at- 
tached a  button-shaped  disk,  the 
face  of  which  is  exposed,  and  is 
on  a  line  with  the  surface  of  the 
anther ;  so  that  these  two  disks 
look  toward  each  other  across  the 
broad  intervening  stigmatic  space, 
as  seen  in  Fig.  461.  The  exposed 
face  of  the  disk  being  covered  with 
a  durable  layer  of  very  viscid  mat- 
ter, the  body  itself  is  sometimes  termed  a  gland,  and  not  improperly. 
The  viscidity  is  nearly  of  the  same  nature  as  that  of  the  interven- 
ing stigma,  of  which  the  glands  are  generally  supposed  to  be 
detached  portions.  If  so,  then  a  portion  of  the  stigma  is  cut  off 
from  the  rest  and  specialized  to  the  purpose  of  conveyance  of  the 
pollen.  When  a  finger's  end  or  any  smaller  body  is  touched  to 
these  disks,  they  adhere  so  firmly  that  the  attached  pollinia  or 
pollen-masses  are  dragged  out  of  the  cell  and  canned  away  en- 
tire. Some  of  these  pollen-masses  have  been  found  attached  by 
the  disk  to  the  eyes  of  a  large  moth.  When  a  moth  of  the  size 
of  head  and  length  of  proboscis  of  Sphynx  drupiferarum  visits  a 
spike  of  these  flowers,  and  presses  its  head  into  the  centre  of  the 


PIG.  463.  A  more  magnified  pollen-mass  of  Platanthera  orbiculata,  with  its  stalk 
and  gland.  464.  Five  of  the  separate  portions  or  pollen-packets,  with  some  of  the 
elastic  threads  of  tissue  connecting  them.  465.  A  portion  more  highly  magnified,  with 
some  of  the  pollen-grains  in  fours  detached. 


ADAPTATIONS   FOR  INTERCROSSING. 


233 


flower  so  that  its  proboscis  may  reach  and  drain  the  bottom  of 
the  nectariferous  tube,  a  pollen-mass  will  usually  be  affixed  to 
each  eye :  on  withdrawal, 
these  will  stand  as  in  Fig. 
466.  Within  a  minute  they 
will  be  turned  downward 
(Fig.  466°),  not  by  their 
weight,  but  by  a  contraction 
in  drying  of  one  side  of  the 
thick  piece  which  connects 
the  disk  with  the  stalk. 
When  a  moth  in  this  con- 
dition passes  from  the  last 
open  flower  of  one  spike  to 
that  of  another  plant,  and 
thrusts  its  proboscis  down  a 
nectary,  the  transported  pol- 
len-masses will  be  brought 
in  contact  with  the  large 
glutinous  stigma :  on  with- 
drawal, either  some  of  the 
small  pellets  of  pollen  will  be 
left  adherent  to  the  stigma, 
the  connecting  elastic  threads 
giving  way  ;  or  else  a  whole 
pollen-mass  will  be  so  left, 
its  adhesion  to  the  glutinous  ^  ° 

stigma  being  greater  than  that  of  the  disk  to  the  moth's  eye. 
The  former  is  a  common  and  a  more  economical  proceeding,  as 
then  a  succession  of  flowers  are  abundantly  fertilized  by  one 
or  two  pollen-masses.  In  either  case,  new  pollen-masses  are 
carried  off  from  fresh  flowers  and  applied  to  the  fertilization 
of  other  blossoms  on  the  same  and  eventually  on  those  of  differ- 
ent individuals.  Cases  like  this,  and  hundreds  more,  all  equally 
remarkable,  serve  to  show  how  sedulous,  sure,  and  economical  are 
the  adaptations  and  processes  of  Nature  for  the  intercrossing  of 
hermaphrodite  flowers. 

42 2a.  An  arrangement  analogous  to  that  of  Orchids,  and 
similarly  subservient  to  cross-fertilization,  characterizes  the 
otherwise  widely  unlike  Asclepias  family.  In  Asclepias  (Milk- 
weed) there  are  five  stamens  surrounding  a  large  stigmatic 

FIG.  466.  Front  part  of  Sphynx  drupiferarum,  bearing  a  pollen-mass  of  Platan- 
thera  orbiculata  affixed  to  each  eye,  in  the  early  position,  466  o.  Front  view  of  the  head, 
later,  showing  the  pollen-masses  deflexed. 


284  THE  FLOWER. 

body,  and  alternating  with  these  five  two-cleft  glands,  the  ver- 
tical chink  or  groove  of  which  is  glutinous.  To  each  gland  is 
firmly  attached,  by  a  caudicle  or  stalk,  a  pollen-mass  of  an  ad- 
jacent anther.  (Fig.  522.)  A  slight  force  upraising  the  gland 
detaches  it  from  the  stigma  and  drags  the  pair  of  suspended 
pollen-masses  out  of  their  cells.  Insects  visiting  the  blossoms 
commonly  dislodge  them,  the  gland  adhering  to  their  legs  or 
tongues  when  these  happen  to  be  drawn  through  the  adhesive 
chink,  and  convey  them  from  one  flower  to  another.  Without 
such  aid  the  flowers  of  Asclepias  rarely  set  seed.1 

423.  Dimorphism,  i.  e.  the  case  of  two  kinds  of  blossoms,  both 
hermaphrodite,  on  the  same  species,  is  another  adaptation  to 
intercrossing.     Not  all  dimorphism,  however,  for  in  cleistogamous 
dimorphism  (434)  the  intent  to  self-fertilize  is  evident.     There 
may  also  be  dimorphism  as  to  the  perianth,  not  particularly 
affecting  fertilization.     One  kind,  however,  and  the  commonest, 
is  a  special  adaptation  to  intercrossing,  viz. : 

424.  Heterogonous  Dimorphism.     (413,  note.)     This  term  is 
applied  to  the  case  in  which  a  species  produces  two  kinds  of 
hermaphrodite  flowers,  occupying  different  individuals,  the  flowers 
essentially  similar  except  in  the  androecium  and  gyncecium,  but 
these  reciprocally  different  in  length  or  height,  and  the  adapta- 
tions such  that,  by  the  agency  of  insects,  the  pollen  from  the 
stamens  of  the  one  sort  reciprocally  fertilizes  the  stigma  of  the 
other.2  This  dimorphism  has  been  detected  in  about  forty  genera 
belonging  to  fourteen  or  fifteen  natural  orders,  widely  scattered 
through  the  vegetable  kingdom  ;  but  there  are  far  more  examples 
among  the  Rubiaceae  than  in  any  other  order.     Sometimes  all 
the  species  of  a  genus  are  heterogonous,  as  in  Houstonia,  and 


1  The  reported  sensitiveness  of  the  gland,  referred  to  in  the  first  issue  of 
this  volume  (1879),  was  founded  upon  misinterpreted  observations. 

*  This  peculiar  arrangement  has  been  long  known  in  a  few  plants,  such 
as  Primula  veris,  P.  grandiflora,  and  Houstonia.  In  Torrey  and  Gray's 
Flora  of  North  America,  ii.  38,  39  (1843),  these  flowers  are  said  to  be  dicecio- 
dimorphous,  not  denoting  that  they  are  at  all  unisexual,  but  that  the  two 
forms  occupy  different  individuals.  Their  meaning  was  detected  by  C. 
Darwin,  and  made  known  in  his  paper  "  On  the  Two  Forms  or  Dimorphic 
Condition  in  the  Species  of  Primula,  and  on  their  Remarkable  Sexual  Rela- 
tions," published  in  the  Journal  of  the  Linnean  Society,  vi.  (1862),  77 :  repub- 
lished,  in  1877,  as  the  leading  chapter  of  his  volume  entitled  "The  Different 
Forms  of  Flowers  on  Plants  of  the  Same  Species."  Mr.  Darwin  had  termed 
these  flowers  simply  Dimorphic;  but  in  this  volume  he  adopted  Hilde- 
brand's  name  of  Heterostyled  for  this  kind  of  blossom.  The  difference, 
however,  affects  the  androecium,  and  even  the  pollen,  as  well  as  the  style ; 
wherefore  we  proposed  for  it  the  name  of  Heterogonous  or  Heterogone  dimor- 
phism, as  mentioned  in  a  former  note,  413. 


ADAPTATIONS   FOE   INTERCROSSING.  235 

Cinchona,  sometimes  only  a  part  of  them,  as  in  Primula  and 
Linum.  In  Hottonia,  a  Primulaceous  genus  of  two  species,  the 
European  one  has  heterogonous  dimorphism *  for  cross-fertiliza- 
tion :  the  American  one  has  homogenous  showy  flowers  with 
only  the  general  chance  for  intercrossing,  and  earlier  flowers 
which  are  cleistogamous  for  self-fertilization. 

425.  The  nature  of  heterogone  dimorphism  may  be  well  under- 
stood from  a  single  example.  The  most  familiar  one  is  that  of 
Houstonia ;  but,  in  larger  blossoms,  Gelsemium  is  a  fine  illus- 
tration in  the  Southern  United  States,  and  Mitchella  (Fig.  467) 
mostly  in  the  Northern.  Raised  from  the  seed,  the  individuals 


are  about  equally  divided  between  the  two  forms  :  namely,  one 
form  with  long  style  and  short  or  low-inserted  stamens ;  the 
other  with  short  style  and  long  or  high-inserted  stamens.  The 
stigmas  in  one  rise  to  about  the  same  height  as  the  stamens  in 
the  other,  both  in  the  tall  or  exserted  organs  and  in  their  low 
and  included  counterparts,  as  is  shown  in  Fig.  468,  answering 
to  the  left  hand  and  Fig.  469  to  the  right  hand  flowers  of  Fig.  467. 
A  bee  or  other  insect  with  proboscis  of  about  the  length  of  the 
corolla-tube,  visiting  the  blossoms  of  Mitchella,  will  brush  the 
same  part  of  its  body  against  the  high  anthers  of  the  long- 
stamened  and  the  high  stigmas  of  the  long-styled  forms ;  and 


1  C.  C.  Sprengel,  as  Darwin  mentions,  had  noticed  this,  before  1793.  He, 
"  with  his  usual  sagacity,  adds  that  he  does  not  believe  the  existence  of 
the  two  forms  to  be  accidental,  though  he  cannot  explain  their  purpose." 
Darwin,  Forms  of  Flowers,  61. 

Some  heterogonous  Primulas  are  said  to  produce  homogonous  varieties  hi 
cultivation.  In  Primula,  and  in  other  genera,  there  are  species  which  seem 
as  if  of  one  sort  only,  no  reciprocal  sort  being  known,  as  if  one  form  had 
become  self-fertile  and  the  other  had  disappeared. 

FIG.  467.  Partridge  Berry,  Mitchella  repens,  In  the  two  forma,  viz.  long-stamened 
and  short-styled,  and  short-stamened  and  long-atyled. 


236  THE   FLOWER. 

the  same  part  of  the  proboscis  against  the  low  anthers  of  the 
short-stamened  and  the  low  stigmas  of  the  short-styled  form. 

426.  Moreover,  Dar- 
win has  ascertained  by 
microscopical  examina- 
tion that  the  pollen  of 
the  two  differs  in  size  or 
shape,  and  by  experi- 
ment that  it  is  less 
active  upon  its  own 
stigma  than  upon  the 
other ;  indeed,  that  in 
many  cases  (as  in  some 
species  of  Linum)  it  is 
quite  inactive  or  impo- 
tent not  only  upon  its 
own  stigma  but  upon  its 
own-form  stigma,  while 
it  is  prepotent  on  the  other,  and  this  reciprocally  of  the  two 
forms.1  Here,  then,  are  flowers  structurally  hermaphrodite,  but 
functionally  as  if  dioecious,  securing  all  the  advantages  of  the 
latter,  along  with  the  economical  advantage  that  both  sorts  of 
individual  and  every  blossom  may  bear  seed.  With  dioecism 
only  about  half  the  plants  could  be  fruitful. 

427.  Heterogonous  Trimorphism.     A   threefold  heterogonism 
is  known  in  certain  species  of  a  few  genera  ;  and  this  complica- 
tion may  have  certain  conceivable  advantages  over  dimorphism. 
Where  seedling  dimorphous  individuals  are  few  and  far  between 
(those  multiplying  from  root  would  all  be  alike),  there  would 
be  an  even  chance  that  any  two  near  each  other  were  of  the 
same  form  and  therefore  sterile  or  imperfectly  fertile.     But  if 
the  organization  were  of  three  forms,  any  two  of  which  inter- 
crossed with  perfect  fertility,  the  chances  (as  Darwin  remarks) 
are  two  to  one  that  any  two  plants  were  of  different  forms,  and 
therefore  by  fertilizing  each  other  completely  fruitful.  • 

428.  The  earliest  known  instance  of  three  forms  as  to  recip> 
rocal  relative  length  of  stamens  and  pistil  is  that  of  Lythrum 

1  Impotence  of  own  pollen,  either  absolute  or  relative,  occurs  no  less  in 
certain  flowers  which  are  not  dimorphous,  as  in  Corydalis,  some  species  of 
Passiflora,  &c.  On  the  contrary,  many  dimorphous  flowers  are  in  a  certain 
degree  self-fertile,  especially  in  the  long-stamened  and  short-styled  form. 
These  subjects  are  physiological,  and  belong  to  another  volume. 

FIG.  468.  Long-styled  flower  of  Pig.  467.  laid  open.  469.  Long-atamened  flower  of 
the  same  laid  open.  Both  equally  enlarged. 


ADAPTATIONS   FOR   INTERCROSSING. 


237 


Salicaria.  This  was  indicated  by  Vaucher  in  1841,  more  par- 
ticularly described  by  Wirtgen  in  1848,  but  was  interpreted  by 
Darwin,  and  the  more  recondite  differences  brought  to  notice,  in 
1864. 1  "  The  three  forms  may  be  conveniently  called,  from  the 
unequal  length  of  their  pistils,  the  long-styled,  mid-styled,  and 
short-styled.  The  stamens  also  are  of  unequal  lengths,  and 
these  may  be  called  the  longest,  mid-length,  and  shortest." 
The  pollen  of  the  different  classes  of  stamens  is  of  two  sorts  as 
to  color,  and  of  three  as  to  size,  the  largest  grains  from  the 
largest  stamens.  "  The  pistil  in  each  form  differs  from  that  in 
either  of  the  other  forms, 
and  in  each  there  are  two 
sets  of  stamens,  different 
in  appearance  and  func- 
tion. But  one  set  of 
stamens  in  each  form 
corresponds  with  a  set 
in  one  of  the  other  two 
forms.  Altogether,  this 
one  species  includes  three 
females  or  female  organs, 
and  three  sets  of  male 
organs,  all  as  distinct 
from  one  another  as  if 
they  belonged  to  different 
species ;  and,  if  smaller 
functional  differences  are 
considered,  there  are  five 
distinct  sets  of  males.  Two  of  the  three  hermaphrodites  mus-w 
coexist,  and  pollen  must  be  carried  by  insects  reciprocally 
from  one  to  the  other,  in  order  that  either  of  the  two  should  be 
fully  fertile  ;  but,  unless  all  three  forms  coexist,  two  sets  of 
stamens  will  be  wasted,  and  the  organization  of  the  species  as 
a  whole  will  be  incomplete.  On  the  other  hand,  when  all  three 
hermaphrodites  coexist,  and  pollen  is  carried  from  one  to  the 
other,  the  scheme  is  perfect :  there  is  no  waste  of  pollen  and  no 

1  In  an  article  On  the  Sexual  Relations  of  the  Three  Forms  of  Lythrum 
Salicaria,  in  Jour.  Linn.  Soc.  viii.  169.  Also  on  the  Character  and  Hybrid- 
like  Nature  of  the  Offspring  of  the  Illegitimate  Unions  of  Dimorphic  and 
Trimorphic  Plants.  Ibid.  x.  393,  1868.  Reproduced  and  extended  in  his 
volume  entitled  "  Forms  of  Flowers,"  1877. 

FIG.  470.  Diagram  of  the  flowers  of  the  three  forms  of  Lythrum  Salicaria,  in  their 
natural  position,  with  the  petals  and  calyx  removed  on  the  near  side.  The  dotted 
lines  with  the  arrows  show  the  directions  in  which  pollen  must  be  carried  to  each 
stigma  to  ensure  full  fertility.  (From  Darwin., 


238  THE  FLOWER. 

false  coadaptation."  The  whole  arrangement  is  displaj'ed  in 
the  annexed  diagram  (Fig.  470) ,  and  in  the  following  account 
of  the  operation.1  "  In  a  state  of  nature,  the  flowers  are  inces- 
santly visited  for  their  nectar  by  hive  and  other  bees,  various 
'Diptera,  and  Lepidoptera.  The  nectar  is  secreted  all  round  the 
base  of  the  ovarium  ;  but  a  passage  is  formed  along  the  upper 
and  inner  side  of  the  flower  by  the  lateral  deflection  (not  repre- 
sented in  the  diagram)  of  the  basal  portions  of  the  filaments ; 
so  that  insects  invariably  alight  on  the  projecting  stamens  and 
pistil  and  insert  their  proboscides  along  the  upper  and  inner 
margin  of  the  corolla.  We  can  now  see  why  the  ends  of  the 
stamens  with  their  anthers  and  the  end  of  the  pistil  with  the 
stigma  are  a  little  upturned,  so  that  they  may  be  brushed  by 
the  lower  hairy  surfaces  of  the  insects'  bodies.  The  shortest 
stamens,  which  lie  enclosed  within  the  calyx  of  the  long-  and 
mid-styled  forms  can  be  touched  only  by  the  proboscis  and  narrow 
chin  of  a  bee :  hence  they  have  their  ends  more  upturned,  and 
they  are  graduated  in  length,  so  as  to  fall  into  a  narrow  file, 
sure  to  be  raked  by  the  thin  intruding  proboscis.  The  anthers 
of  the  longer  stamens  stand  laterally  farther  apart  and  are  more 
nearly  on  the  same  level,  for  they  have  to  brush  against  the 
whole  breadth  of  the  insect's  body.  .  .  Now  I  have  found  no 
exception  to  the  rule  that,  when  the  stamens  and  pistil  are  bent, 
they  bend  to  that  side  of  the  flower  which  secretes  nectar.  .  .  . 
When  nectar  is  secreted  on  all  sides,  they  bend  to  that  side 
where  the  structure  of  the  flower  allows  the  easiest  access  to  it, 
as  in  Lythrum.  ...  In  each  of  the  three  forms,  two  sets  of  sta- 
mens correspond  in  length  with  the  pistil  in  the  other  two  forms. 
When  bees  suck  the  flowers,  the  anthers  of  the  longest  stamens, 
bearing  the  green  pollen,  are  rubbed  against  the  abdomen  and 
the  inner  sides  of  the  hind  legs,  as  is  likewise  the  stigma  of  the 
long-styled  form.  The  anthers  of  the  mid-length  stamens  and 
the  stigma  of  the  mid-styled  form  are  rubbed  against  the  under 
side  of  the  thorax  and  between  the  front  pair  of  legs.  And, 
lastly,  the  anthers  of  the  shortest  stamens  and  the  stigma  of  the 
short-styled  form  are  rubbed  against  the  proboscis  and  chin  ;  for 
the  bees  in  sucking  the  flowers  insert  only  the  front  part  of  their 
heads  into  the  flower.  On  catching  bees,  I  observed  much  green 
pollen  on  the  inner  sides  of  the  hind  legs  and  on  the  abdomen, 
and  much  yellow  pollen  on  the  under  side  of  the  thorax.  There 
was  also  pollen  on  the  chin,  and,  it  may  be  presumed,  on  the 
proboscis,  but  this  was  difficult  to  observe.  I  had,  however, 

*  All  from  Darwin,  Forms  of  Flowers,  137-147,  &c. 


ADAPTATIONS  FOB  INTERCROSSING.  23iJ 

independent  proof  that  pollen  is  carried  on  the  proboscis  ;  for  a 
small  branch  of  a  protected  short-styled  plant  (which  produced 
spontaneously  only  two  capsules)  was  accidentally  left  during 
several  days  pressing  against  the  net,  and  bees  were  seen  insert- 
ing their  proboscides  through  the  meshes,  and  in  consequence 
numerous  capsules  were  formed  on  this  one  small  branch.  .  .  . 
It  must  not,  however,  be  supposed  that  the  bees  do  not  get  more 
or  less  dusted  all  over  with  the  several  kinds  of  pollen ;  for  this 
could  be  seen  to  occur  with  the  green  pollen  from  the  longest 
stamens.  .  .  .  Hence  insects,  and  chiefly  bees,  act  both  as 
general  carriers  of  pollen,  and  as  special  carriers  of  the  right 
sort." 

429.  Finally,  a  long  series  of  experiments  (requiring  eighteen 
distinct  kinds  of  union)  proved  that  both  kinds  of  pollen  are 
nearly  or  quite  impotent  upon  the  stigma  of  the  same  flower,  and 
that  no  ovary  is  fully  fertilizable  by  other  than  a  "  legitimate 
union,"  i.  e.  by  stamens  of  the  corresponding  length ;  but  'that 
the  mid-length  pistil  is  more  prolific  than  either  of  the  others 
under  illegitimate  union  of  either  kind ;  which  might  perhaps  be 
expected,  as  the  pollen  proper  to  it  is  intermediate  in  size  of 
grains  between  that  of  the  long  and  that  of  the  shortest  stamens. 

430.  Nesaea  verticillata,  a  common  Lythraceous  plant  of  the 
Atlantic  United  States,  is  similarly  trimorphous,   but  has  not 
yet  been  particularly  investigated.     Several  South  African  and 
American  species  of  Oxalis  are  equally  trimorphous,  and  have 
been   investigated  by  Darwin   and   Hildebrand,1  with  results 
quite  as   decisive  as  in  Lythrum   Salicaria.      One  genus  of 
Monocotyledons  has  trimorphous  blossoms,  viz.  Pontederia,  of 
which  the  North  American  P.  cordata  is  a  good  illustration.8 

431.  All  known  flowers  exhibiting  reciprocal  dimorphism  or 
trimorphism  are   entomophilous  :  no  such  wind-fertilized  species 
is  known.     Few  of  them  are  irregular,  and  none  very  irregular : 
they  do  not  occur,   for  instance,   in  Leguminosae,   Labiatae, 

1  Monatsber.  Akad.  Berlin,  1866 ;  Bot.  Zeit.  1871,  &c.    According  to 
Darwin,  Fritz  Mueller  "  has  seen  in  Brazil  a  large  field,  many  acres  in  extent, 
covered  with  the  red  blossoms  of  one  form  [of  an  Oxalis]  alone,  and  these 
did  not  produce  a  single  seed.     His  own  land  is  covered  with  the  short-styled 
form  of  another  species,  and  this  is  equally  sterile;  but,  when  the  three 
forms  were  planted  near  together  in  his  garden,  they  seeded  freely."  Forms 
of  Flowers,  180. 

2  Detected  by  W.  H.  Leggett.     See  Bulletin  of  Torrey  Bot.  Club,  vi.  62, 
170;  and  for  the  original  discovery  in  Brazilian  species,  by  Fritz  Mueller, 
see  Darwin's  Forms  of  Flowers,  183,  &c.     Pontederia  has  three  lengths  of 
style  and  counterpart  stamens,  as  in  Lythrum  Salicaria,  each  flower  having 
two  sets  of  stamens.,  three  in  each  set. 


240  THE   FLOWER. 

Scrophulariacese,  Orchidaceae,  &c.  Nature  is  not  prodigal,  and 
does  not  endow  with  needless  adaptations  flowers  which  are 
otherwise  provided  for. 

§  3.  ADAPTATIONS  FOR  CLOSE  FERTILIZATION. 

432.  Even   where   cross-fertilization   in   bisexual  flowers   is 
obviously  arranged  for,  it  is  apt  to  be  tempered  with  more  or 
less  of  close-fertilization.     The  more  exquisite  the  arrangements 
for  the  former  are,  the  more  completely  is  the  plant  dependent 
upon  insect  visitation.     Failure  to  intercross  is  a  remote  and 
small  evil  compared  with  failure  to  set  seed  at  all.     In  order 
therefore  that  the  plan  of  cross-fertilization  may  not  defeat  even 
its  own  end,  through  too  absolute  dependence  on  precarious 
assistance,  some  opportunity  for  self-fertilization  will  usually  be 
advantageous.     Also   there   is   a   long   array   of  insect-visited 
flowers,  especially  polyandrous  ones,  in  which  close  fertilization 
must  be  much  the  commoner  result,  except  where  the  pollen  of 
another  but  wholly  similar  flower  has  greater  potency. 

433.  Subsidiary  self-fertilization  is  secured  in  a  great  variety 
of  ways.     In  Gentiana  Andrewsii,  which  is  proterandrous,  and 
usually  cross-fertilized  by  humble-bees  entering  bodily  into  the 
corolla,  an  exposed  surface  of  pollen  long  remains  fresh  upon 
the  ring  of  anthers  girding  the  base  of  the  style  :  when  the  stigmas 
separate,  they  remain  for  some  days  simply  divergent,  but  they 
at  length  become  so  revolute  that  the  receptive  surface  is  brought 
into  contact  with  the  ring  of  pollen  below.     The  opening  and 
closing  of  blossoms  by  day  or  night,  the  growth  of  style,  fila- 
ments, or  corolla  after  anthesis  commences,  or  other  changes  of 
position,  may  secure  a  certain  amount  of  self-fertilization  in  a 
subsidiary  or  even  in  a  regular  way.     Then  certain  species,  such 
as  Chickweed,   which   blossom   through  a  long  season,  close- 
fertilize  even  in  the  bud  in  early  spring,  when  insects  are  scarce, 
but  are  habitually  intercrossed  by  insects  in  summer.     Somewhat 
similarly,  according  to  Hermann  Mueller,1  certain  species,  such 
as  Euphrasia  officinalis  and  Rhinanthus  Crista-galli,  habitually 
produce  two  kinds  of  blossoms,  one  larger  and  more  showy, 
usually  affecting  sunny  localities,   and  with  parts  adapted  to 
intercrossing  by  insects  ;  the  other  smaller  or  inconspicuous,  and 
with  anthers  adjusted  for  giving  pollen  to  the  adjacent  stigma 
without  aid.     There  are  gradations  between  these  last  arrange- 
ments, and  the  more  special  and  remarkable  one  of  dimorphism 
with 

1  Befruchtung  der  Blumen  durch  Insekten,  294 ;  Nature,  viii.  433. 


ADAPTATIONS   FOR,   CLOSE   FERTILIZATION.          241 

434.  Cleistogamy.     Here  the  intention  and  the  accomplishment 
of  self-fertilization  are  unmistakable.     This  peculiar  dimorphism 
consists  in  the  production  of  very  small  or  inconspicuous  and 
closed   flowers,   necessarily  self-fertilized   and   fully   fertile,  in 
addition  to  ordinary,  conspicuous,  and  much  less  fertile,  though 
perfect  flowers.     Two  cases  were  known  to  Linnaeus,1  and  one 
of  them  to  Dillenius  before  him  ;  those  of  Viola  have  long  been 
familiar  in  the  acaulescent  species ;  Adrien  Jussieu  made  out 
the  structure  of  the  cleistogamous  flowers  in  certain  Malpighiaceae 
in  1832,  and  recorded  in  1843  that  Adolphe  Brongniart  had  well 
investigated  those  of  Specularia,  and  that  Weddel  had  discov- 
ered them  in  Impatiens  Nolitangere.     A  full  account  of  the  then 
known  cases  was  given  by  Mohl 2  in  1863  ;  but  D.  Mueller,  of 
Upsala,  who  examined  Viola  canina,  is  said  by  Darwin  to  have 
given,8  in  1857,  "the  first  full  and  satisfactory  account  of  any 
cleistogamic  flower."     The   appropriate   name  of  cleistogamous 
was  given  by  Kuhn,4  in  1867,  and  is  now  in  common  use. 

435.  Cleistogamous  flowers  are  now  known  in  about  60  genera, 
of  between  twenty  and  thirty  natural  orders,  of  very  various 
relationship,  though  all  but  five  are  Dicotyledons.     All  but  the 
Grasses5  and  Juncus  are   entomophilous   as   to   the   ordinary 
flowers,  and  most  of  these  such  as  have  special  arrangements  for 
their  intercrossing,  either  by  dichogamy,  heterogone  dimorphism 
or  trimorphism  (in  Oxalis) ,  or  such  special  contrivances  as  those 
of  Orchids. 

436.  It  has  been  said  that  the  ordinary  flowers  in  such  plants 
are  sterile,  and  perhaps  they  always  are  so  except  when  cross- 
fertilized  :  in  most  cases  they  are  habitually  infertile  or  spar- 
ingly fertile.     Probably  they   suffice   to   secure   in   every   few 
generations  such  benefit  as  a  cross  may  give,  while  the  principal 


1  Campanula  (now  Specularia)  perfoliata  and  Euellia  clandestine  the 
latter  a  cleistogamous  state  of  R.  tuberosa.  Linnseus  did  not  make  out  the 
structure  of  the  flowers,  but  supposed  them  to  want  the  stamens. 

*  In  Bot.  Zeitung,  xxi.  309. 
8  In  Bot.  Zeitung,  xvi.  730. 

*  Ibid.  xxv.  65.  The  name  ( denoting  "  closed  up  "  union  or  fertilization )  has 
been  written  deistogenous,  which  is  not  so  proper.     We  prefer  cleistogamous  to 
cleistogamic  (and  so  of  similar  terms),  as  best  harmonizing  with  the  Latin 
adjective  form,  both  in  form  of  termination  and  in  euphoniously  taking  the 
accent  upon  the  antepenult. 

8  Amphicarpum  (Milium  amphicarpon,  Pursh)  is  the  earliest  recognized 
cleistogamous  Grass,  except  perhaps  Leersia  oryzoides.  Some  species  of 
Sporobolus  are  like  the  latter,  and  Mr.  C.  G.  Pringle  has  recently  detected 
such  flowers  concealed  at  the  base  of  the  sheaths  in  Danthonia.  Amer. 
Jour.  Sci.  January,  1878,  71. 

19 


242  THE  FLOWER. 

increase  is  by  cleistogamous  self-fertilization,  which  thus  offsets 
the  incidental  disadvantage  of  the  former  mode. 

437.  In  general,  the  cleistogamous  are  like  unto  the  ordinary 
flowers  arrested  in  development,  some  arrested  in  the  almost 
fully  formed  bud,  most  at  an  earlier  stage,   and  in  the  best 
marked   cases   with    considerable   adaptive    modification.      In 
these,  "their  petals  are  rudimentary  or  quite  aborted;   their 
stamens  are  often  reduced  in  number,  with  anthers  of  very  small 
size,  containing  few  pollen-grains,  which  have  remarkably  thin 
transparent   coats,  and  generally  emit  their  tubes  while  still 
enclosed  within  the  anther-cells  ;  and,  lastly,  the  pistil  is  much 
reduced  in  size,  with  the  stigma  in  some  cases  hardly  at  all 
developed.     These  flowers  do  not  secrete  nectar  or  emit  any 
odor :  from  their  small  size,  as  well  as  from  the  corolla  being 
rudimentary,  they  are  singularly  inconspicuous.     Consequently, 
insects  do  not  visit  them ;  nor,  if  they  did,  could  they  find  an 
entrance.     Such  flowers  are  therefore  invariably  self-fertilized  ; 
yet  they  produce  an  abundance  of  seed.     In  several  cases,  the 
young  capsules  bury  themselves  beneath  the  ground,  and  the 
seeds  are  there  matured.     These  flowers  are  developed  before, 
or  after,  or  simultaneously  with  the  perfect  ones."  *     In  Grasses, 
however,  as  in  some  Dicotyledons,  there  is  much  less  modifica- 
tion and  more  transition.     For  when  Leersia  half  protrudes  its 
panicle,  in  the  usual  way,  the  included  half  is  fertile  and  the 
expanded  portion  sterile  (or  almost  always  so),  although  the 
flowers  may  open  and  exhibit  well-developed  anthers,  ovaries, 
and  stigmas.     But  when  similar  panicles  remain  enclosed  in  the 
leaf-sheaths,  they  are  mostly  fruitful  throughout. 

438.  Fully  to  apprehend  the  economy  of  cleistogamy  in  pollen- 
saving   alone,  —  and  contrariwise  to  estimate   the  expense  of 
intercrossing,  —  one  should  compare  the  small  number  of  pollen- 
grains  which  so  completely  serve  the  purpose  in  a  typical  cleis- 
togamous flower  (say  400  in  Oxalis  Acetosella,  250  in  Impatiens, 
100  in  some  Violets)  with  the  several  thousands  of  all  entomo- 
philous  cross-fertilized  flowers,  rising  to  over  three  and  a  half 
millions  in  the  flower  of  a  Peony,  also  their  still  greater  number 
in  manj^  anemophilous  blossoms.     To  this  loss  should  be  added 
the  cost  of  a  corolla  and  its  action,  also  of  the  production  of 
odorous  material  and  of  nectar.     No  species  is  altogether  cleis- 
togamous.    Thus  cleistogamy,  with  all  its  special  advantage, 
testifies  to  the  value  of  intercrossing. 


1  Darwin,  Forms  of  Flowers,  310. 


PERIANTH,  OR   FLOWER-LEAVES.  243 

SECTION  V.     THE  PERIANTH,1  OB  CALYX  AND  COROLLA  ra 

PARTICULAR. 

439.  The  distribution  of  the  floral  leaves   around  the  axis, 
which  belongs  to  phyllotaxy,  and  their  particular  disposition  in 
the  bud  (aestivation) ,  have  already  been  considered  in  Chap.  IV. 
Sect.  I.,  II.     And  most  of  the  morphology  of  calyx  and  corolla 
has  been  outlined  in  the  preceding  sections  of  the  present  chap- 
ter.    What  remains  chiefly  relates  to  particulars  of  form  and 
to  terminology. 

440.  Duration.     The  differences  in  this  respect  give  rise  to  a 
few  terms,  such  as  the  following.     Calyx  or  corolla  may  be 

Persistent,  not  cast  off  after  anthesis,  but  remaining  unwithered 
until  the  fruit  is  formed  or  matured ;  as  the  calyx  in  Labiatae,  in 
Physalis,  and  most  Roses. 

Marcescent,  withering  or  drying  without  falling  away ;  as  the 
corolla  of  Heaths,  Drosera,  &c. 

Deciduous,  falling  after  anthesis  and  before  fructification  ;  as 
the  petals  of  Roses,  the  calyx  and  corolla  of  Columbine. 

Ephemeral  or  Fugacious,  lasting  for  onlv  a  day  ;  as  the  petals 
of  Poppy,  Helianthemum,  Purslane,  and  Spiderwort.  In  the 
two  former,  they  are  cast  or  early  deciduous,  the  anthesis  lasting 
but  a  day :  in  the  two  latter,  the  anthesis  is  equally  or  more 
brief,  but  the  petals  deliquesce  or  decay  at  once  without  falling, 
as  does  the  whole  flower  of  Cereus  grandiflorus  and  other  night- 
blooming  Cactaceae. 

Caducous,  falling  when  the  blossom  opens ;  as  the  calyx  of 
Poppy  and  Baneberry. 

441.  Numerical   Terms,   succinctly  denoting  the   number   of 
leaves,  either  of  the  perianth  as  a  whole,  or  of  any  one  of  its 
circles,  are  common  in  descriptive  botany.     The  most  general 
are  those  which  simply  specify  the  number  of  component  leaves, 
by  prefixing  Greek  numerals  to  the  Greek  name  of  leaves,  ex- 
pressing them  in  Latin  form,  or  transferring  them  to  the  Eng- 
lish.    Thus 

Diphyllous,  of  two  leaves  (sepals  or  petals)  ;  Triphyllous,  of 
three  ;  Tetraphyllous,offour  ;  Pentaphyllous,  of  five  ;  ffexaphyllous, 
of  six,  and  so  on.  A  tulip  and  a  Tradescantia  flower  have  a 
hexaphyllous  perianth,  but  composed  of  two  circles,  answering 
to  calyx  and  corolla ;  each  Triphyllous*  When  the  character 

1  Perianthium,  alias  Perigone  or  Periganium.  (296.) 

2  As  elsewhere  explained,  when  numerical  composition  is  indicated  without 
reference  to  nature  of  parts,  the  terms  dimerous,  trimerous,  tetramerous,  penta 
merous,  &c.,  may  be  used.  (322.) 


244  THE  FLOWEB. 

of  the  organ,  i.  e.  whether  calyx  or  corolla  is  to  be  specified, 
the  word  sepal  or  petal  is  employed  in  the  combination ;  as, 

Disepalous,  of  two  sepals  ;  Trisepalous,  of  three  ;  Tetrasepalous, 
of  four ;  Pentasepalous,  of  five  (also  written  5-sepalous,  and  ac- 
cordingly 2-sepalous,  3-sepalous) ,  and  so  on  :  also, 

Dipetalous,  Tripetcdous,  Tetrapetalous,  Pentapetalous  (2-5- 
petalous),  &c.,  when  the  corolla  is  concerned. 

442.  Monophyttous,    Mbnosepalous,   and  Monopetalous  are  the 
proper  terms  for  perianth    (calyx,  corolla,  &c.)   composed  of  a 
single  leaf.     Likewise  Polyphyllous,  Polysepalous,  and  Polypetal- 
ous  for  the  case  of  a  considerable  but  unspecified  number  of 
members.     Unfortunately,  in   the  Linnsean  and  long-prevalent 
use,  monopetalous  was  the  term  employed  to  designate  a  corolla 
of  one  piece  in  the  sense,  or  the  fact,  of  a  coalescence  or  grow- 
ing together  of  two,  three,  five,  or  more  petals  into  a  cup  or 
tube ;  and  so  of  a  calyx,  of  a  whorl  of  bracts,  &c.     And  poly- 
petalous,  polysepalous,  and  polyphyllous  were  the  counterparts 
of  this,  meaning  of  more  than  one  distinct  piece,  whatever  the 
number.     The  misleading  use,  consecrated  by  long  prescription, 
is  not  yet  abandoned,  but  will  in  time  be  obsolete.     In  present 
descriptive   botany,    a  polyphyllous   calyx,  or   a   poly  petalous 
corolla,  or  a  5 -petalous  corolla,  would  be  taken  to  mean  that  the 
sepals  or  petals  (as  the  case  may  be)  were  distinct  or  uncom- 
bined,  and  a  monopetalous  corolla  to  be  one  with  petals  combined 
by  coalescence.  (329.) 

443.  Terms  of  Union  or  Separation.     The  proper  term  for  a 
corolla  or  a  calyx  the  leaves  of  which  are  more  or  less  coalescent 
into  a  cup  or  tube  is 

Gamopetalous  for  such  a  corolla,  Gamosepalous  for  the  calyx  ; 
these  terms  meaning  united  petals  or  sepals.  The  older  and  mis- 
leading names  Monopetalous  and  Monosepalous,  although  current 
up  to  a  recent  day,  should  be  discontinued.  Another  term  is 
not  rarely  used  in  Germany,  that  of  Sympetalous,  for  the  gamo- 
petalous  (or  formerly  monopetalous)  corolla,  —  therefore  Syn- 
sepalous  for  a  similar  calyx.  It  is  perhaps  a  more  apt  term 
than  gamopetalous,  and  of  the  same  etymological  signification  ; 
but  the  latter  is  already  well  in  use. 

Choripetalous  is,  on  the  whole,  the  most  fitting  name  for  a 
corolla  the  petals  of  which  are  separate  (as  it  literally  expresses 
this),  that  is,  for  what  is  still  commonly  called  Polypetalous,  as 
already  explained.  (442.)  It  is  adopted  by  Eichler,  &c.  Chori- 
sepalous  is  the  term  applied  to  the  calyx.  Dialypetcdous  (em- 
ployed by  Endlicher)  has  the  same  meaning.  Both  this  term 
and  choripetalous  carry  the  implication  of  separated,  rather 


PERIANTH,  OB  FLOWER-LEAVES.  245 

than   of  typically  separate,  parts.      Eleutheropetalous  (literally 
free-petalled)  has  also  been  used,  but  is  inconveniently  long. 

444.  Degree  of  coalescence  is  most  correctly  expressed  by  the 
phrases  united  (connate,  or  coherent,  or  coalescent)  at  the  base, 
to  the  middle,  or  to  the  summit,  as  the  case  may  be.     But  it  is 
more  usually  and  tersely  expressed  in  botanical  description  by 
employing  terms  of  division,  identical  with  those  used  in  describ- 
ing  the  lobing   or  toothing   of  leaves   and   all  plane   organs. 
(184-188.)     That  is,  the  calyx  or  corolla  when  gamophyllous 
is  for  description  taken  as  a  whole,  and  is  said  to  be  parted 
(3-parted,  b-parted,  &c.),  when  the  sinuses   extend  almost  to 
the  base  ;  cleft,  when  about  to  the  middle  ;  lobed,  a  general  term 
for  any  considerable   separation  beyond   toothing ;    dentate  or 
toothed    (%-toothed,    b-toothed,    &c.),   when   the    union   extends 
almost  to  the  summit ;  entire,  when  the  union  is  complete  to 
the  summit  or  border. 

445.  Parts  of  Petals,  &c.     The  expanded  portion  of  a  petal, 
like  that  of  a  leaf,  is  the  LAMINA  or  BLADE  :  any  much  contracted 
base  is  the  UNGUIS  or  CLAW.     The  latter  is  very  short  in  a  rose- 


petal,  but  long  and  conspicuous  in  a  pink  and  all  flowers  of  that 
tribe  (Fig.  471),  in  many  Capparidese  (Fig.  409)  and  Cruciferae. 
A  sepal  is  very  rarely  distinguishable  into  lamina  and  claw. 

446.  Parts  of  Gamophyllous  Perianth.  The  coalescent  portion 
of  a  corolla,  calyx,  or  of  a  perianth  composed  of  both  (such  as 
a  Lily  or  Crocus-blossom) ,  so  far  as  the  sides  are  parallel  or  not 
too  spreading,  is  its  TUBE  :  an  expanded  terminal  portion,  either 
divided  or  undivided,  is  the  LIMB  or  BORDER.  The  limb  may 


FIG.  471.  Corolla  of  Soapwort,  of  five  separate  long-clawed  or  unguiculate  petals, 
with  a  crown  at  the  junction  of  claw  and  blade. 

FIG.  472.  Flower  of  Gilia  coronopifolia ;  the  parts  answering  to  the  claws  of  the 
petals  of  the  last  figure  here  all  united  into  a  tube. 

FIG.  473.  Flower  of  the  Cypress-Vine  (Ipomcea  Quamoclit) ;  the  petals  a  little  farther 
united  into  a  flve-lobed  spreading  border. 

FIG.  474.  Flower  of  the  Ipomoea  coccinea ;  the  five  component  petals  perfectly  united 
into  a  trumpet-shaped  tube,  and  beyond  Into  an  almost  entire  spreading  border. 


246  THE  FLOWER. 

be  parted  (that  is,  the  component  parts  not  united)  quite  or 
nearly  down  to  the  tube  or  base,  as  in  Fig.  472,  475  ;  or  less  so, 
as  in  Fig.  473,  476  (with  limb 
5-lobed)  ;  or  with  merely  angles 
or  points  to  represent  the  tips 
of  the  component  members,  as  in 
Fig.  474  ;  or  with  even  and  entire 
border,  as  in  common  Morning- 
Glory,  Fig.  482. 

447.  The  line,  or  sometimes  a  manifest  or  conspicuous  portion, 
between  the  limb  and  tube  (in  the  corolla  always  a  portion  above 
or  at  the  insertion  of  the  stamens,  when  these  are  borne  by  the 
corolla)  is  called  the  THROAT,  in  Latin  FAUX,  pi.  fauces.     This 
is  mostly  more  open  than  the  tube,  yet  less  expanded  than  the 
limb  ;  but  it  often  presents  insensible  gradations  from  the  one  to 
the  other. 

448.  Such  appendages  as  the  CORONA  or  CROWN  (385,  shown 
in  Fig.  403,  404,  471)  usually  belong  to  the  throat  of  a  gamo- 
petalous  corolla  or  perianth,  as  in  Oleander,  Comfrey,  Borrage, 
Narcissus,  &c.,  or  to  a  corresponding  position  when  the  parts 
are  not  coalescent. 

449.  Forms  of  Corolla,  Calyx,  &c.     As  to  terminology,  some 
of  these  are  special  and  are  applicable  to  corolla  only,  as  the 

Papilionaceous,  the  peculiar  irregular  corolla  of  the  typical 
portion  of  Leguminosse  (388,  Fig.  342-334),  which  has  been 
already  illustrated,  and  in  which  the  petals,  two  pairs  and  an 
odd  one,  take  particular  names.  Also  the 

Caryophyllaceous,  or  Pink-flower  (Fig.  471),  a  regular  corolla, 
of  five  long-clawed  (unguiculate)  petals,  the  claws  enclosed  in  a 
tubular  calyx  and  the  blades  spreading  ;  and  the 

Cruciferous,  of  four  somewhat  similar  petals,  the  four  abruptly 
spreading  blades  in  the  form  of  a  cross  (cruciate),  as  in  Fig.  394. 

Rosaceous,  with  roundish  and  widely  spreading  petals  on  very 
short  or  hardly  any  claws,  as  in  Rose  and  Apple-blossoms. 

Liliaceous,  a  6-phyllous  perianth  of  campanulate  or  funnelform 
shape  ;  the  members  either  distinct,  as  in  most  common  lilies  and 
tulips,  or  gamophyllous,  as  in  Lily  of  the  Valley.  All  but  the 
first  and  last  of  these  sorts  are  examples  of  regular  and  chori- 
petalous  perianth. 

Orchidaceous  flowers  are  of  a  peculiar  irregularity,  combining 
both  calyx  and  corolla :  one  member,  the  petal  in  front  of  the 

FIQ.  475.  Rotate  or  wheel-shaped  and  five-parted  corolla  of  the  Bittersweet  (So- 
lanuni  Dulcamara). 

FIQ.  476.    Wheel-shaped  and  five-lobed  corolla  of  the  common  Potato. 


PERIANTH,  OR   FLOWER-LEAVES.  247 

stamen  and  stigma,  differs  from  the  rest  in  shape  and  in  being 
nectariferous  (as  in  Fig.  460)  ;  it  is  named  the  LABELLUM. 

Galeate  is  a  term  applied  to  a  corolla  the  upper  petal  or  part 
of  waich  is  arched  into  the  shape  of  a  casque  or  helmet,  called 
the  Galea;  as  in  Aconite  (Fig.  357)  and  Lamium,  Fig.  479. 
In  the  former  the  galea  is  of  a  single  petal ;  in  the  latter,  it 
consists  of  two,  completely  united. 

450.  Gamophyllous  forms  with  special  names  are  chiefly  the 
following.  Illustrations  are  usually  taken  from  the  corolla,  but 
the  forms  and  terms  are  not  peculiar  to  it,  excepting  the  first, 
viz.  the 

Ligulate  or  Strap-shaped  corolla  (Fig.  288,  &c.),  which  is 
nearly  confined  to  Compositae.  Here  a  corolla,  formed  of  three 
or  five  petals,  imitates  a  single  petal,  except  at  its  very  base, 
which  is  commonly  tubular :  the  remainder  is  as  though  the  tube 
had  been  split  down  on  the  upper  side  and  flattened  out.  The 
corolla  of  Lobelia,  t}^pe  of  a  family  most  nearly  related  to  Com- 
positae, illustrates  this.  (Fig.  488.) 


451.  The  names  of  the  general  forms  are  mostly  taken  from 
some  resemblance  to  common  objects.  All  those  in  common 
use  will  be  found  in  the  Glossary :  a  few  leading  ones  are  here 
specified.  They  may  be  divided  into  the  regular  and  the  irregu- 
lar. The  principal  irregular  form  with  a  special  name  is  the 

Labiate,  or  lipped,  also  termed  BilaUate,  as  there  are  two  lips, 
an  upper  and  a  lower  (superior  and  inferior,  or  anterior  and 
posterior,  290),  although  one  of  them  is  sometimes  obscure  or 
abortive.  This  bilabiate  character  in  the  corolla,  and  often  in 
the  calyx  also,  pervades  several  orders  with  gamopetalous 
flowers,  and  gives  name  to  one  of  them,  the  Labiatas,  to  which 

FIG.  477.  Campanulate  corolla  of  the  Harebell,  Campanula  rotundifolia.  478. 
Salverform  (hypocraterimorphous)  corolla  of  Phlox.  479.  Labiate  (ringent)  corolla  of 
Lamium ;  a  side  view  480.  Personate  corolla  of  Antirrhinum  or  Snapdragon.  481. 
Personate  corolla  of  Linaria,  spurred  (calcarate)  at  the  base. 


248  THE  FLOWER. 

the  Sage  and  Mint  belong.  Such  flowers  are  5-merous,  and 
have  two  members  specially  united  to  form  one  lip,  and  three  in 
the  other.  The  odd  sepal  being  posterior  (or  next  the  axis  of 
inflorescence) ,  and  consequently  the  odd  petal  anterior,  the  calyx 
has  its  lower  lip  of  two  sepals  and  its  upper  of  three  ;  while  the 
corolla  has  its  upper  lip  of  two  petals  and  its  lower  of  three. 
But  in  Leguminosae,  where  the  calyx  is  sometimes  bilabiate,  and 
where  the  odd  sepal  is  anterior  (or  toward  the  bract),  this  is 
reversed,  and  two  sepals  or  lobes  of  the  calyx  form  the  upper 
lip  and  three  the  lower.  A  bilabiate  corolla  is 

Ringent,  that  is  gaping  or  open-mouthed,  when  the  throat  is 
freely  open,  as  in  Lamium,  Fig.  479  ; 

Personate,  or  masked,  when  the  throat  is  closed,  more  or  less, 
by  a  projection  of  the  lower  lip  called  the  PALATE,  as  in  Antir- 
rhinum and  Linaria,  Fig.  480,  481. 

452.  Of  regular  forms,  there  are  the  following,  beginning  with 
that  having  least  tube : 

Rotate,  or  Wheel-shaped  (Fig.  475,  476),  widely  spreading  from 
the  very  base,  or  from  a  short  and  inconspicuous  tube. 

Crateriform,  or  Saucer-shaped,  like  rotate  except  that  the  broad 
limb  is  cupped  by  some  upturning  toward  the  margin. 

Hypocrateriform,  or  rather  (not  to  mix  Latin  and  Greek) 
Hypocraterimorphous ,  in  English  Salverform,  when  a  rotate  or 
saucer-shaped  limb  is  raised  on  a 
slender  tube  which  does  not  much 
enlarge  upward  ;  that  is,  where  a  long 
and  narrow  tube  abruptly  expands 
into  a  flat  or  flattish  limb,  as  in 
Fig.  478.  In  Fig.  472-474  are  seen 
salverform  corollas  with  somewhat 
more  upwardly  dilated  (trumpet- 
shaped)  tube.  The  salver  or  hypo- 
craterium,  which  the  name  refers  to, 
with  a  stem  or  handle  beneath,  is  now 
to  be  met  with  only  in  old  pictures. 

Tubular,  when  strictly  used,  denotes 
a  gamophyllous  perianth  with  limb 
inconspicuous  in  proportion  to  the 
tube,  as  in  Trumpet  Honeysuckle,  or  as  Fig.  472-474  would  be 
if  the  limb  were  much  diminished  or  wanting.  But  it  is  some- 
times used  in  the  sense  of  having  a  conspicuous  tube. 


FIG.  482.    Calyx  and  funnelform  (infundibuliform)  corolla  of  a  common  Morning- 
Glory,  Ipomnea  purpurea. 


ANDKCECIUM,   OR  STAMENS. 


249 


Infundibuliform,  or  Funnelform,  such  as  the  corolla  of  common 
Morning-Glory  (Fig.  482) ,  denotes  a  tube  gradually  enlarging 
upward  from  a  narrow  base  into  an  expanding  border  or  limb. 

Campanulate,  or  Bell-shaped  (Fig.  477),  denotes  a  tube  of  length 
not  more  than  twice  the  breadth,  moderately  expanded  almost 
from  the  base,  the  sides  above  little  divergent. 


SECTION  VI.    THE  ANDKOECIUM,  OR  STAMENS  IN  PARTICULAR. 

453.  The  whole  Stamen.  For  the  general  character  and  some 
of  the  modifications  of  the  stamens,  see  the  first  (301)  and  por- 
tions of  the  succeeding  sections  of  the  present  chapter.  The 
terms  peculiar  to  these  organs,  and  of  common  use  in  botanical 
description,  were  nearly  all  coined  by  Linnaeus,  and  employed  as 
the  names  of  classes  in  his  sexual  system.  (672.)  The  sub- 
stantive names  of  those  classes  which  are  characterized  by  the 
number  of  stamens,  and  which  were  designated  by  Greek  nume- 
rals prefixed  to  andria  (the  Greek  word  for  man  being  used 
metaphorically  for  stamen),  are  put  into  adjective  form,  as 
follows : 

Monandrous,  for  a  flower  with  a  solitary  stamen ;  Diandrous, 
for  a  flower  with  two  stamens  ;  Triandroits,  with  three  ;  Tetran- 
drous,  with  four  ;  Pentandrous,  with  five  ;  Hexandrous,  with  six  ; 
Heptandrous,  with  seven  ;  Octandrous,  with  eight ;  Enneandrous, 
with  nine ;  Decandrous,  with  ten ;  Dodecandrous,  with  twelve ; 


Polyandrous,  with  a  greater  or  indefinite  number,  or  Icosandrous 
(meaning  twenty-stamened)  when  a  polyandrous  flower  has  the 
stamens  inserted  on  the  calyx,  as  in  the  Cherry  (Fig.  337), 
Pear,  &c. 

FTG.  483.    Diadelphous  stamens  (9  and  1)  of  a  Pea.    484.  Monadelphous  stamens  of 
aT.npine.    485.  Monadelphous  stamens,  &c.,  of  Mallow. 

FIG.  486.  Five  syngenesious  stamens  of  a  Composita.    487.  The  same,  laid  open. 


250 


THE   FLOWER. 


Didynamous  is  a  term  applied  to  an  androecium  of  four  sta- 
mens in  two  pairs,  a  longer  and  a  shorter,  as  in  Fig.  361. 

Tetradynamous  is  similarly  applied  to  that  of  six  stamens,  two  of 
them  shorter,  in  the  manner  characteristic  of  Cruciferse,  Fig.  396. 

454.  Terms  which  denote  coalescence  of  stamens,  whether  by 
their  filaments  or  their  anthers,  are 

Monadelphous,  that  is,  in  one  brotherhood,  by  coalescence  of 
the  filaments  into  a  tube,  as  in  the  Mallow  (Fig.  485) ,  Lupine 
(Fig.  484),  Lobelia  (Fig.  488),  &c. 

Diadelphous,  in  two  brotherhoods,  by  coalescence  of  the  fila- 
ments into  two  sets  ;  sometimes  an  equal  number  in  each,  as  in 
Fumariaceae  (Fig.  390),  sometimes  nine  in  one  set  and  one 
separate,  as  in  the  Pea  (Fig.  483)  and  most  Papilionaceae. 

Triadelphous,  with  filaments  united  in  three  sets  or  clusters,  as 
in  Hypericum. 

Pentadelphous,  in  five  sets,  as  in  Linden,  Fig.  398,  399. 
But  in  general,  when  the  sets  are  several,  without  regard  to  the 
number  the  stamens  are  said  to  be  Polyadelphous. 

Syngenesious,  when  the  stamens  are  united  by  their  anthers 
into  a  tube  or  ring ;  as  in  the  whole  vast  order  of  Composite 


(Fig.  486,  where  they  are  five  in  number  and  the  filaments  dis- 
tinct), in  Cucurbita  (Fig.  489,  490,  where  they  are  three  in 
number  and  the  filaments  partly  monadelphous) ,  and  in  Lobelia 
(Fig.  488,  where  they  are  also  five  and  the  long  filaments  are 
mainly  monadelphous) . 

FIG.  488.  Flower  of  Lobelia  cardinalis,  with  tube  of  corolla  divided  on  one  side ; 
filaments  and  anthers  united  into  a  tube:  /.  tube  of  filaments;  a.  of  anthers. 

FIG.  489.  Male  flower  of  Cucurbita  (Squash),  with  limb  of  calyx  and  corolla  cut 
away,  to  show  the  stamens,  viz.,  three  filaments,  separate  at  base  but  monadelphous 
above,  and  three  syngenesious  anthers  in  a  kind  of  head.  490.  Stamens  of  the  same, 
enlarged  and  the  upper  part  cut  away,  to  show  the  union.  The  anthers  are  sinuous. 
491.  A  detached  stamen  of  the  Melon,  with  loosely  sinuous  anther. 

Fig.  492.  Stamens  and  style  of  a  Cypripedium,  united  into  one  body  or  column: 
o.  anthers;  it.  enlarged  sterile  stamen;  stig.  the  stigma. 


ANDRCECIUM,  OR   STAMENS.  251 

455.  Of  terms  relating  to  adnation  of  stamens,  besides  the 
general  ones   of  hypogynous,  perigynous,  epigynous   (332),   and 
epipetalous,  or  adnate  with  corolla,  there  is  the  special  one  of 

Gynandrous,  having  stamens  borne  upon  the  pistil,  as  in 
Orchidacese.  In  C}rpripedium,  the  filaments  of  two  stamens,  and 
an  enlarged  sterile  stamen  behind,  are  adnate  to  a  style,  while 
the  two  anthers  are  quite  free  (Fig.  492)  ;  in  the  proper  Orchis 
tribe  (as  in  Fig.  460,  461),  anther  and  stigma  are  consolidated 
into  one  mass,  and  there  is  no  evident  style. 

456.  A  complete  stamen  consists  of  FILAMENT  and  ANTHER. 
The  latter  is  the  functionally  essential  part  of  the  organ,  and 
therefore  is  wanting  only  in  abortive  or  sterile  stamens.  (345, 
352,  &c.)     The  filament,  being  only  a  stalk  or  support,  may  be 
very  short  or  wholly  wanting :  then  the  anther  is  sessile,  just  as 
the  blade  of  a  leaf  is  said  to  be  sessile  when  there  is  no  petiole. 

457.  The  Filament,   although  usually  slender  and  stalk-like, 
assumes  a  great  variety  of  forms  :  it  is  sometimes  dilated  so  as 
to  resemble  a  petal,  except  by  its  bearing  an  anther ;  as  in  the 
transition  states  between  the  true  petals  and  stamens  of  Nym- 
phsea,  shown  in  Fig.  318. 

458.  Such  petaloid  filaments  would  indicate  that  this  part  of 
the  stamen  answered  to  blade  rather  than  to  footstalk,  while 
others  would  harmonize  better  with  what  seems  at  first  sight  to 
be  the  more  natural  view,  that  the  filament  is  the  homologue  of 
the  petiole,  the  anther  of  the  blade  of  a  leaf.     Eemembering 
that  in  large  numbers  of  leaves  there  is  no  distinction  into  petiole 
and  lamina  or  blade,  such  homologies  should  not  be  insisted  on. 
The  filament  may  be  variously  appendaged  by  outgrowths.    Some 
of  these  appendages  are  very  conspicuous,  such  as  the  scale  of 
Larrea  (Fig.  405) ,  which  is  on  the  inside,  and  the  nectariferous 
hood  of  Asclepias  on  the  outside ;  or  there  may  be  a  tooth  on 
each  margin,  as  in  species  of  Allium. 

459.  The  Anther,  the  essential  organ  of  the  stamen,  contain- 
ing the  pollen,  surmounts  the  filament,  when  that  is  present. 
It  normally  consists  of  two  cells  or  lobes,  the  word  cell  being 
here  used  in  the  sense  of  sac.     But,  as  each  sac  is  not  rarely 
divided  into  two  cavities  (locelli),  the  best  technical  name  for 
anther-sac  is  that  of  THECA.     The  two  thecae,  lobes,  or  cells  are 
commonly  connected  by  a  more  or  less  evident  and  sometimes 
conspicuous  common  base  or  junction,  which  is  mostly  a  pro- 
longation  of    the   filament,    the   CONNECTIVUM,  or  in   English 
CONNECTIVE. 

460.  For  the  discharge  of  the  pollen,  the  cells  of  a  normal 
anther  open  at  the   proper  time   by  a  line  or  chink,  usually 


252 


THE  FLOWER. 


extending  from  top  to  bottom  (Fig.  493),  the  suture  or  line  of 
dehiscence.  Commonly  this  line  is  lateral  or  marginal :  not 
rarely  it  faces  forward  or  backward.  In  the  vast  genus  Solanum, 


to  which  the  Potato  belongs,  in  most  Ericaceous  plants  (Fig. 
458,  494),  in  Polygala,  and  in  many  other  flowers,  the  anther- 
cells  open  only  by  a  hole  (foramen  or  pore) ,  or  at  most  a  short 
chink,  at  the  tip,  through  which  the  pollen  has  in 
some  way  to  be  discharged.  In  Vaccinium  (Cran- 
berry, Blueberry,  &c.),  the  pore-bearing  tip  of  the 
anther-cell  is  prolonged  considerably,  often  into  a 
slender  tube,  as  in  Fig.  340.  In  the  Barberry  (Fig. 
495)  and  in  most  of  that  family,  also  in  Lauraceae, 
the  whole  face  of  each  anther-cell  separates  by  a  con- 
tinuous line,  forming  a  kind  of  door,  which  is  attached  at  the  top, 
and  turns  back,  as  if  on  a  hinge  :  in  this  case,  the  anthers  are  said 
to  open  by  uplifted  valves.  In  the  Sassafras  and  many  other 
plants  of  the  Laurel  family,  each  lobe  of  the  anther  opens  by  two 
smaller  valves  of  the  kind,  like  trap-doors. 

461.  The  attachment  of  the  anther  to  the  filament  presents 
three  principal  modes,  which  are  connected  by  gradations. 
These  are  the 

Innate  (Fig.  495,  496),  in  which  the  anther  directly  continues 
and  corresponds  to  the  apex  of  the  filament,  the  cells  usually 
dehiscent  strictly  marginally,  the  lobes  or  cells  not  looking  or 
projecting  either  inward  or  outward. 

FIG.  493.  A  stamen,  with  its  anther,  &,  surmounting  the  filament,  a,  and  opening 
in  the  normal  manner  down  the  whole  length  of  the  outer  side  of  each  cell. 

FIG.  494.  Stamen  of  a  Pyrola ;  each  cell  of  the  anther  opening  by  a  terminal  ori- 
fice or  pore. 

FIG.  495.  Stamen  of  a  Barberry;  the  cells  of  the  anther  opening  each  by  an  up- 
lifted valve. 

FIG.  496.  A  stamen  of  Isopyrum  biternatum,  with  innate  anther.  497.  Stamen  of 
Liriodendron,  or  Tulip-tree,  with  adnate  extrorse  anther.  498.  Stamen  of  OSnothera 
glauca,  with  the  anther  fixed  by  its  middle  and  versatile. 

FIG.  499.  A  stamen  of  Asarum  Canadense,  with  adnate  anther  and  prolonged  tip 
to  connective. 


ANDRCECIUM,  OR   STAMENS.  253 

Adnate,  in  which  the  connective  appears  to  be  a  direct  con- 
tinuation of  the  filament,  having  the  anther  adherent  to  the 
anterior  or  posterior  face  of  it,  and  the  lines  of  dehiscence 
therefore  looking  inward  or  outward.  Magnolia,  Liriodendron 
(Fig.  497),  and  Asarum  (Fig.  499)  furnish  good  examples  ;  the 
latter  conspicuously  so,  on  account  of  a  prominent  prolongation 
of  the  connective  or  tip  of  the  filament. 

Versatile,  when  the  anther  is  attached  at  some  part  only  of  its 
back  or  front  to  the  tip  of  the  filament,  on  which  in  anthesis  it 
lightly  swings  ;  as  in  Plantain,  in  all  Grasses,  the  Lily,  Evening 
Primrose  (Fig.  498) ,  &c. 

462.  The  direction  to  which  an  anther  faces,  whether  inward 
(toward  the  centre  of  the  flower),  or  outward  (toward  the  peri- 
anth) ,  has  to  be  considered ;  except  in  the  case  of  an  innate 
anther  with  strictly  lateral  or  marginal  dehiscence.    An  anther  is 

Extrorse,  i.  e.  turned  outward,  or  Posticous,  when  it  faces  to- 
ward the  perianth,  as  in  Magnolia  and  Liriodendron  (Fig.  497), 
Asarum  (Fig.  499),  and  Iris ;  these  all  being  cases  of  adnate 
and  extrorse  anthers,  the  cells  attached  for  their  whole  length  to 
the  outside  of  the  summit  of  the  filament  or  the  connective. 

Introrse,  i.  e.  turned  inward,  or  Anticous,  when  it  faces  toward 
the  axis  of  the  flower ;  as  in  Nymphaeacese  (Fig.  318),  in  Violet 
and  Lobelia  (which  are  adnate  and  introrse) ,  and  in  GEnothera. 
In  the  common  Evening  Primroses  (as  in  Fig.  498)  the  anther 
is  fixed  near  the  middle,  introrse,  and  versatile. 

463.  The  direction  in  which  the  anther  may  be  said  to  face, 
outward  or  inward,  depends  upon  two  characters,  which  do  not 
always  coincide,  viz.  the  insertion  or  attachment  of  the  cells, 
and  the  position  of  their  line  of  dehiscence.     In  such  a  strongly 
characterized  adnate  anther  as  that  of  Liriodendron  (Fig.  497), 
both  the  attachment  and  the  dehiscence  are  plainly  posticous  or 
extrorse :  in  most  species  of  Trillium,  the  cells  are  introrse  as  to 
attachment,  but  some  are  nearly  marginal  and  some  are  even 
rather  extrorse  as  to  dehiscence  :  in  the  related  Medeola,  and  in 
Lilium,  where  the  anthers  are  extrorsely  affixed  toward  the  base 
or  middle  to  a  slender  tip  of  the  filament,  the  dehiscence  is 
either  introrse  or  nearly  marginal.     Parnassia  is  in  similar  case  ; 
the  anthers  being  clearly  extrorse  as  to  insertion  and  more  or 
less  introrse  as  to  dehiscence. 

464.  Adnate  anthers  are   perhaps  as  frequently  extrorse  as 
introrse.     Others,  whether  basifixed  or  medifixed,  are  more  com- 
monly introrse.      Those  fixed  by  the  middle,  or  at  any  other 
part  of  the  back,  and  lying  on  the  inner  side  of  the  filament, 
are  said  to  be  Incumbent. 


254  THE  FLOWER. 

465.  The  connective  may  be  appendaged  either  by  a  prolon- 
gation or  otherwise  from  the  tip  (as  in  Fig.  499),  or  from  the 
back,  as  in  Violets  and  in  many  Ericaceous  plants. 

466.  The  normal  anther  is  two-celled,  bilocidar,  or  (to  use 
a  less  common  term)  dithecous,  and  its  lobes  or  cells  parallel, 
right  and  left ;  but  the  cells  at  first,  and  sometimes  at  maturity, 
are  bilocellate,  that  is  each  is  divided  into  two  by  a  partition 
which  stretches   from  the  connective  to  the  suture  or  line  of 

dehiscence.  In  an 
innate  anther,  and 
in  many  others,  this 
line  of  dehiscence  is 
marginal  or  lateral, 
either  strictly  or 

«»  "an"  '  ste'         nearly     so,     as     in 

Fig.  500.  When  introrse  or  extrorse  (as  in  Fig.  501,  502), 
the  sutures  ma}7  still  be  considered  to  represent  the  margins 
turned  inward  or  outward.  The  pollen  is  accordingly  pro- 
duced in  four  cavities  or  separate  portions  of  the  interior.  But 
ihe  two  locelli  on  the  same  side  of  the  midrib  or  connective 
(right  and  left)  are  usually  confluent  into  one  pollen-filled  cavity 
or  cell  at  maturity  if  not  earlier,  or  at  least  the  partition  between 
them  breaks  up  at  dehiscence.  Sometimes  it  remains,  and,  the 
groove  at  the  sutures  being  deep,  the  anther  is  strongly  four- 
lobed  or  quadrilocular  at  maturity,  as  in  Menispermum  (Fig. 
504)  ;  but  morphologically  this  is  still  only  bilocular  (dithecous) 
although  quadrilocellate,  and  the  anther  opens  at  the  sutures 
and  through  these  partitions. 

467.  A  stamen  being  the  homologue  of  a  leaf,  the  natural 
supposition  is  that  the  anther  is  homologous  with  the  blade  or 
an  apical  portion  of  the  blade,  therefore  the  two  lobes  or  thecae 
with  the  right  and  left  halves  of  it,  the  intervening  connective 
with  th'e  midrib,  and  the  line  of  dehiscence  with  the  leaf-mar- 
gins.1   This   conception  is   exemplified  by  the  accompanying 

1  This  is  the  view  long  ago  taken  by  Cassini  and  Keeper,  and  it  may  still 
be  maintained  as  the  best  morphological  conception.  Mohl  interposed  some 
objections  to  its  universality ;  but,  as  presented  in  Sachs's  Text-Book,  they 
are  not  incompatible  with  the  common  morphology.  Sachs  takes  the  fila- 
ment with  the  connective  to  be  the  homologue  of  the  whole  leaf,  and  the 
anther-cells  as  appendages.  Others,  in  likening  the  anthers  to  glands,  adopt 
a  similar  view. 

FIG.  500.  Innate  anther,  same  as  Fig.  496,  in  younger  state,  with  transverse  section, 
•bowing  the  four  locelli.  501.  Same  of  an  adnate  extrorse  anther,  such  as  Fig.  497. 
502.  Same  as  the  preceding  but  mature  and  dehiscent,  the  two  locelli  becoming  one  cell 
by  the  vanishing  or  breaking  up  of  the  partition. 


ANDRCECIUM,  OB    STAMENS. 


255 


diagram,  Fig.  503,  which  should,  however,   show  the  median 

partitions  in  the  cross-section,  or  traces  of  them.     Pollen  is  a 

special  development  into  peculiar  cells  of  what  would 

be  parenchyma  in  a  leaf.      Its  formation  normally 

begins  in  four  places,  which  may  remain  separate 

up  to  maturity,  or  the  two  on  each  side  of  the  axis 

or  connective  may  early  be  confluent  into  one  cell. 

468.  Of  the  many  deviations  of  the  typical  two- 
celled   anther,   with   its   cells   parallel  and   united 
longitudinally  by  a  connective,  the   simplest  and 
commonest  is  that  in  which  (as  in  Fig.  505)  the 
two  cells  diverge  below  and  remain  united  only  at 
their  apex.     Next,  the  two  cells  may,  in  their  early 
development,  become  confluent  at  the  apex,  as  in  the 

Mallow  family  (Fig.  506),  so  as  to  form  a  continuous  pollin- 
iferous  cavity  within,  opening  by  a  continuous  suture  round  the 
margin :  here  the  anther  is  unilocular  or  one-celled  by  confluence. 
In  another  way,  the  anthers  of  some  species  of  Orthocarpus  (gen- 
erally resembling  Fig.  505,  but  the  lobes  or  cells  quite  distinct 
or  even  separated  at  apex)  lose  one  of  the  cells  by  partial  or 
complete  non-development  and  so  become  one-celled  by  abor- 
tion. The  anther  of  Gomphrena  (Fig.  507)  is  completely  uni- 
locular by  abortion  or  suppression  of  the  companion  cell.  Thus 
losing  one  half,  it  is  said  to  be  dimidiate,  or  halved. 

469.  The  two  anther-cells,  such  as  those  of  Fig.  505,  some- 
times diverge  so  much  that  they  form  a  straight  line  transverse 
to  the  filament,  as  in  Monarda  (Fig. 

508),  in  which  their  contiguous  ends 
so  coalesce  as  to  give  the  appearance 
of  a  one-celled  anther  fixed  by  the 
middle.  Or,  again,  the  two  cells  may 
be  separated  by  the  enlargement  of 
the  connective  between  them,  as  in  Cal- 
amintha,  Fig.  509.  This  enlargement 
is  extreme  in  the  great  genus  Salvia,  in 
which  a  very  long  and  narrow  connec- 
tive gives  the  appearance  of  a  filament  astride  the  apex  of  the 

FIG.  603.  Diagram  to  illustrate  the  morphology  of  the  stamen,  on  the  idea  that  the 
anther  answers  to  leaf-blade:  the  lower  portion  being  filament  and  a  part  of  the  anther, 
in  section,  the  upper  a  part  of  a  leaf. 

FIG.  504.    Stamen  of  Menispermum  Canadense,  the  quadrilocellate  anther  divided. 

FIG.  605.    Stamen  of  Pentstemon  pubescens,  with  anther-cells  divergent. 

FIG.  506.  Stamen  of  Mallow  (one  of  the  cluster  of  Fig.  485),  the  two  cells  and  •utures 
confluent  into  one. 

FIG.  507.  Anther  of  Gomphrena  or  Globe  Amaranth,  mediated,  of  a  single  cell, 
dehiscent. 


256 


THE   FLOWEB. 


proper  filament,  and  bearing  an  anther-cell  at  each  end.  In  a 
few  species,  the  two  anther-cells  are  nearly  alike  ;  in  more,  the 
lower  one  is  imperfect,  as  in  Fig.  510" ;  in  more,  it  is  abortive 
or  wanting  altogether,  as  in  Fig.  510*.  Then,  in  the  related 
Californian  genus  Audibertia,  the  lower  half  of  this  connective 


fs  reduced  to  a  short  tail,  as  shown  in  Fig.  511°,  or  even  in 
most  of  the  species  to  so  minute  a  vestige  that,  except  for  these 
transitions,  the  stamen  might  be  supposed  to  consist  of  a  simple 
filament,  with  an  interruption  like  a  splice  in  the  middle,  and 
surmounted  by  a  one-celled  anther,  as  shown  in  Fig.  511*.  In 
Rosemary,  the  continuity  is  complete,  although  a  minute  reflexed 
tooth  sometimes  indicates  the  junction. 

470.  Pollen,  the  product  of  the  anther,  is  usually  a  powdery 
substance,  which  when  magnified  is  seen  to  consist  of  separate 
grains,  of  definite  size  and  shape,  uniform  in  the  same  plant, 


but  often  very  different  in  different  species  or  families.     The 
grains  are  commonly  single  cells,  globular  or  oval  in  shape,  and  • 
of  a  yellow  color.     But  in  Spiderwort  they  are  oblong ;  in  the 

FIG.  508-511.  Anthers,  with  upper  part  of  filament,  of  several  Labiatse.  608.  Of 
Monarda.  509.  Of  a  Calamintha.  510  Of  two  species  of  Sal  via,  with  long  and  slender 
connective,  the  upper  fork  of  which  bears  one  anther-cell ;  the  lower  in  a  (from  Salvia 
Texana),  bearing  the  other  cell  in  an  imperfect  condition ;  in  b  (from  S.  coccinea),  bear- 
Ing  none  at  all.  511.  a.  Same  of  Audibertia  grandiflora,  the  lower  fork  of  the  connec- 
tive reduced  to  a  naked  spur ;  b,  from  A.  stachyoides,  in  which  this  lower  fork  is  nearly 
wanting,  and  the  upper  is  in  a  straight  line  with  the  filament  which  it  seems  to 
continue. 

FIG.  512-515.  Forms  of  pollen:  512,  from  Mlmulus  mowhatus;  513,  Sicyos;  614, 
Echinocystis;  515,  Hibiscus. 


POLLEN.  257 

Cichory  and  Thistle  tribes,   many-sided;  in  the  Musk-plant, 
spirally  grooved ;    in  the  Mallow  family  and  the  Squash  and 


Pumpkin,  beset  with  briskly  projections,  &c.     The  pollen  of 

Pine,  as  well  as  that  of  the  Onagraceae,  is  not  so  simple,  but 

appears  to  consist  of  three  or  four  blended  cells  ; 

that  of  most  Ericaceae  evidently  consists  of  four 

grains  or  cells  united.   (Fig.  512-521.)      The 

most  extraordinary  shape  is  that  of  Zostera,  or 

the  Eel-grass  of  salt-water,  in  which  the  grains 

(destitute  of  the  outer  coat)  consist  of  Jong  and 

slender  threads,  which,  as  they  lie  side  by  side  in  the  anther, 

resemble  a  skein  of  silk. 

471.  Pollen-grains  are  usually  formed  in  fours,  by  the  division 
of  the  living  contents  of  mother  cells  first  into  two,  and  these 
again  into  two  parts,  which  become  specialized  cells.     As  the 
pollen  completes  its  growth,  the  walls  of  the  mother  cells  are 
usually  obliterated.      But   sometimes   these   cells 

persist,  either  as  shreds,  forming  the  cobweb-like 
threads  mixed  with  the  pollen  of  Evening  Primrose, 
or  as  a  kind  of  tissue  combining  the  pollen  into 
coherent  masses,  of  various  consistence.  Of  this 
kind  are  the  elastically  coherent  pollen-masses  (or 
POLLINIA,  sing.  POLLINIUM)  of  Orchises  (Fig.  463) ,  m 

and  the  denser  waxy  ones  of  many  other  orchids  and  those  of 
Asclepias  or  Milkweed,  Fig.  522. 

472.  A  pollen-grain  has  two  coats.     The  outer  coat  is  com- 
paratively thick,  and  often  granular  or  fleshy.     This  is  later 
formed  than  the  inner,  and  by  a  kind  of  secretion  from  it :  to  it 
all  the  markings  belong.     The  inner  coat,  which  is  the  proper 
cell- wall,  is  a  very  thin,  delicate,  transparent  and  colorless  mem- 
brane, of  considerable  strength  for  its  thickness.     The  pollen 
of  Zostera  and  of  some  other  aquatic  plants  is  destitute  of  the 
outer  coat. 

473.  The  cavity  enclosed  by  the  coats  is  filled  with  a  viscid 
substance,  which  often  appears  slightly  turbid  under  the  higher 
powers  of  ordinary  microscopes,  and,  when  submitted  to  a  mag- 

PfG.  516-521.    Forms  of  pollen:  516,  Lily;  517,  Cichory;  518,  Pine;  519,  Circsea; 
520.  Kalmia;  521,  Evening  Primrose. 

FIG.  522.    A  pair  of  pollinia  of  Asclepias,  annexed  by  their  caudicles  to  the  gland. 
17 


258  THE  FLOWER. 

nifying  power  of  about  three  hundred  diameters,  is  found  to 
contain  a  multitude  of  minute  particles  (fovittce) ,  the  larger  of 
which  are  from  one  four-thousandth  to  one  five-thousandth  of  an 
inch  in  length,  and  the  smaller  only  one  fourth  or  one  sixth  of 
this  size.  When  wetted,  the  grains  of  pollen  promptly  imbibe 
water  by  endosmosis,  and  are  distended,  changing  their  shape 
somewhat,  and  obliterating  the  longitudinal  folds,  one  or  more 
in  number,  which  many  grains  exhibit  in  the  dry  state.  Soon 
the  more  extensible  and  elastic  inner  coat  inclines  to  force  its 
way  through  the  weaker  parts  of  the  outer,  especially  at  one  or 
more  thin  points  or  pores ;  sometimes  forming  projections, 
when  the  absorption  is  slow  and  the  exterior  coating  tough.  In 
many  kinds  of  pollen,  the  grains,  when  immersed  in  water,  soon 
distend  to  bursting,  discharging  the  contents.1 

474.  Pollen- tubes.     In  others,  and  in  most  fresh  pollen,  when 
placed  in  ordinarily  aerated  water,  at  least  when  this  is  slightly 
thickened  by  sj'rup  or  the  like,  and  submitted  to  a  congenial  tenv 
perature,  a  projection  of  the  inner  coat  through  the  outer  appears 
at  some  one  point,  and  by  a  kind  of  germination  grows  into  a 
slender  tube,  which  may  even  attain  two  or  three  hundred  times 
the  diameter  of  the  grain  ;  and  the  richer  protoplasmic  contents 
tend  to  accumulate  at  the  farther  and  somewhat  enlarging  ex- 
tremity of  this  pollen-tube.2 

475.  In  cleistogamous  flowers  (434),  the  pollen,  while  still  in 
the  anther,  sends  out  its  tubes,  which  may  grow  to  a  great  length, 
in  the  mere  moisture  of  the  flower-bud,  the  growing  tip  always 
directing  itself  toward  the  stigma  in  a  wonderful  way.     Similarly, 
in  the  open  flower  of  Milkweeds,  the  pollen-tubes  sometimes 
start  from  the  pollen-mass  even  while  yet  in  the  anther,  and  in 
vast  numbers,  forming  a  tuft  or  skein  of  pollen-tubes,  which 
may  attain  considerable  length  and  direct  itself  toward  the  some- 
what distant  stigma.     Commonly,  however,  the  pollen  remains 

1  In  Coniferae,  the  grams  of  pollen  have  a  peculiar  internal  structure  or 
rather  a  development  (suggestive  of  a  homology  with  the  microspores  of  some 
of  the  higher  Cryptogamia),  the  contents  at  or  before  maturity  undergoing 
division  into  two  or  three  internal  cells,  only  one  of  which  acts  in  fertiliza- 
tion. When  they  act  upon  the  ovule  or  are  placed  in  water,  and  the  inner 
coat  swells  by  absorption,  the  bursting  outer  coat  is  commonly  thrown  off. 
In  Pines  and  Firs  (but  not  in  Larch  and  Hemlock  Spruce),  the  grain  of 
pollen  is  singularly  compound,  consisting  (as  in  Fig.  518)  of  a  central  arcuate 
body  (the  proper  pollen-cell)  bearing  at  each  end  an  empty  roundish  cell. 
These  are  vesicular  protrusions  or  appendages  of  the  proper  pollen-grain,  of 
no  known  functional  importance,  except  that  they  render  such  wind-dis- 
persed pollen  more  buoyant  for  transportation. 

a  Van  Tieghem,  in  Ann.  Sci.  Nat.  ser.  6,  xii.  312,  &c.,  1869. 


GYNCECIUM  IN  ANGIOSPEKMS.  259 

unaltered  until  it  is  placed  upon  the  stigma.  The  more  or  less 
viscid  moisture  of  this  incites  a  sim- 
ilar growth,  and  also  doubtless  nour- 
ishes it ;  and  the  protruding  tube  at 
once  penetrates  the  stigma,  and  by  glid- 
ing between  its  loose  cells  buries  itself 
in  the  tissue  of  the  style,  descending 
thence  to  the  interior  of  the  ovary  and 
at  length  to  the  ovules.  Fertilization 
is  accomplished  by  the  action  of  this 
pollen-tube  upon  the  ovule,  and  upon  a 
special  formation  within  it.  Consequent 
upon  this  an  embryo  is  formed ;  and  the  ovule  now  becomes  a  seed. 


SECTION  VII.     THE  PISTILS,  OR  GYNCECTUM. 
§  1.    Ix  ANGIOSPERMS. 

476.  The  succinct  description  of  the  pistil  in  the  first  section 
of  this  chapter  (302),  as  also  what  has  been  stated  of  the  modi- 
fications of  the  gyncecium  in  Section  III.,  relates  to  the  most 
typical  conditions  of  this  part   of  the   flower.     The   essential 
characteristics  of  all  ordinary  pistils,  whether  simple  or  compound, 
are  :  1 .  a  closed  ovary,  in  which  one  or  more  ovules  are  included  ; 
and  2.  a  stigma,  upon  which  pollen  for  fertilizing  the  ovules  is 
received,  and  through  which  the  pollen  acts  upon  them.     There 
is  a  more  simplified  condition,  in  Gymnosperms,  in  which  naked 
ovules  are  exposed  to  the  direct  action  of  the  pollen.     In  con- 
tradistinction to  this,  the  ordinary  pistil  is  said  to  be  Angiosper- 
mous ;  that  is,  with  the  seeds  enclosed  in  a  sac  or  covering,  this 
in  the  flower  being  the  ovary.1    And  plants  with  such  gynoecium 
are  denominated  ANGIOSPERMS  or  ANGIOSPERMOUS  PLANTS.     To 
such  only  the  present  subsection  specifically  relates. 

477.  The   several  terms  which   apply  to   the    Gynacium   or 
female  system  of  a  flower,  and  to  its  components,  have  been 

1  Although  thus  originated,  the  seeds  are  not  in  all  cases  matured  in  a 
closed  pistil.  In  the  Blue  Cohosh,  Caulophyllum  thalictroides,  the  ovules 
rupture  the  ovary  soon  after  flowering,  and  the  seeds  become  naked ;  and  in 
Mignonette  they  are  imperfectly  enclosed,  the  ovary  being  open  at  the 
summit  from  an  early  period  of  fructification. 

FIG.  523.  A  pollen-grain  of  Datura  Stramonium,  emitting  its  tube.  524.  Pollen- 
grain  of  a  Convolvulus,  with  its  tube.  525.  Other  pollen-grains,  with  their  tubes,  less 
strongly  magnified.  526  A  pollen-grain  of  the  Evening  Primrose,  resting  on  a  portion 
of  the  stigma,  into  which  the  tube  emitted  from  one  of  the  angles  penetrates;  the  oppo- 
site angle  also  emitting  a  pollen-tube.  All  highly  magnified. 


260 


THE  FLOWER. 


enumerated  and  defined  already  (302,  note)  :  the  elementary 
term  is  that  of 

478.  Carpel,  Lat.  CARPELLUM.   This  is  the  term  coined  by  Dunal, 
and  is  in  common  use.      The  better-  formed  word  CARPIDIUM 
(English   Carpid)  has  been  proposed,  and  best  of  all  CARPO- 
PHYLLDM,  in  English  Carpophyll.     For  carpels  are,  as  the  word 
carpophylla  denotes,  pistil-  leaves,  or  leaves  of  the  gyncecium, 
t.  e.,  seed-bearing  or  fructiferous  phylla.     They  occupy  the  cen- 
tral or  uppermost  region  of  the  flower.     A  carpel  may  be  a  pistil 
of  itself,  either  the  only  one  of  a  blossom  or  one  of  several,  or 
it  may  be  a  constituent  of  a  more  complex  pistil.     In  either  case, 
a  carpel  is  the  homologue  of  a  leaf. 

479.  The  morphological  conception  of  an  uncombined  carpel 
is  that   of  the  blade  of  a  leaf  incurved  lengthwise,   so   that 
the  margins  meet,  and  join  by  a  suture,  thus  forming  a  closed 
sac,  the  ovary.     A  prolongation  of  the  tip  of  the  leaf  is  the  style: 
some  portion  of  this,  usually  the  apex,  not  rarely  a  single  or 
double  line  down  the  side  which  answers  to  the  suture  of  the 
leaf-margins,  and  may  be  regarded  as  its  continuation,  is  the 
stigma.     The  carpellary  leaf  is  always  wcurved  :  the  lower  sur- 

face of  the  leaf  is  represented  by 
the  exterior  surface  of  the  ovary, 
the  upper  by  the  interior.  The 
conjoined  margins  of  the  leaf,  or 
whatever  they  bear,  are  internal 
in  the  ovary  :  the  stigma  may  be 
regarded  as  a  portion  of  leaf- 
margins  presented  externally,  des- 
titute of  epidermis  and  formed 
of  loose  cellular  tissue,  which  in 
anthesis  is  moist  by  some  secretion.  The  ovules  are  peculiar 
structures  normally  arising  as  outgrowths  from  the  margins  of 
the  leaf,  or  some  part  of  them,  sometimes  from  the  whole  or 
a  special  portion  of  the  upper  or  inner  surface  of  the  leaf. 

480.  The  carpellary  leaf  being  involute,  the  suture,  on  which 
the  ovules  are  normally  borne,  always  looks  toward  the  axis  or 
centre  of  the  flower.     It  is  the  only  proper  suture  (or  seam)  a 
carpel  can  have.     From  its  position  it  takes  the  name  of  Inner 
or  Ventral  Suture.     And  the  opposite  line  or  ridge,  answering 
to  the  midrib  of  the  leaf,  being  sometimes  prominent  and  of  the 

FIG.  527.  A  leaf  incurving,  to  illustrate  the  morphology  of  a  simple  pistil  or  carpel. 
628.  A  carpel  (of  Isopyrum  biternatum),  cut  aoross,  the  lateral  stigma  (here  manifestly 
a  double  line)  and  the  suture  bearing  the  ovnles  tnrne<l  t^wnrd  the  eye.  529.  A  ripe 
carpel  of  ^'ars'i  Marigold  which  has  opened  and  shed  the  seeds:  the  points  of  attach- 
ment of  the  latter  conspicuous  along  the  edges  of  the  carpel. 


GYNCECITJM   IN   ANGIOSPEBMS.  261 

appearance  of  a  suture,  has  been  somewhat  incongruously  named 
the  Outer  or  Dorsal  Suture. 

481 .  The  number  of  carpels  in  a  gynoecium  is  simply  expressed 
by  adjective  terms  consisting  of  Greek  numerals  prefixed  to  this 
word  :  e.  g.,  Monocarpellary,  of  a  solitary  carpel ;  Dicarpellary,  of 
two  carpels  ;   Tricarpellary,  of  three  ;   Tetracarpellary,  of  four ; 
Pentacarpellary ,  of  five,  and  so  on  up  to  Polycarpettary,  of  many 
or  at  least  of  several  and  an  indefinite  number.     Less  general 
and  only  partially  synonymous  terms  are  such  as  Monogynous 
(of  one  pistil) ,  Digynous  (of  two) ,  Polygynous  (of  many) ,  &c. 
These  are  adjective  forms  of  the  names  of  the  orders,  from 
Monogynia  to  Polygynia,  in  the  Linnaean  artificial  classification, 
which  either  supposes  the  carpels  to  be  separate  or  partly  so,  or 
confounds  simple  and  compound  pistils. 

482.  When  the  gynoacium  is  of  a  solitary  carpel,  the  position 
of  this  as  regards  the  axis  of  inflorescence  is  not  uniform ;  but 
commonly  its  back  or  dorsal  suture  is  before  the  subtending 
bract,  or  in  other  words  the  ventral  or  ovule-bearing  suture 
faces  the  axis  of  inflorescence.     When  there  are  two  carpels, 
they  face  each  other,  bringing  their  ventral  sutures  into  opposi- 
tion, and  as  to  axis  of  inflorescence  either  median  or  transverse 
(291),  but  usually  median,  that  is  antero-posterior  or  in  the 
line  of  bract  and  axis.     Cruciferae,  Capparidaceae,  and  Fumari- 
aceae  are  somewhat  remarkable  for  having  their  two  carpels 
right  and  left,  that  is,  collateral  or,  in  other  words,  transverse. 
When  three,  four,  or  a  greater  number,  they  divide  the  circle 
equally,  or  when  numerous  they  take  a  spiral  instead  of  verticil- 
late  order,  and  occupy  several  or  many  ranks,  as  in  Ranunculus, 
Magnolia,  Potentilla,  &c. 

483.  The    Gynoecium    may  be    either    of   separate  carpels 
(Apocarpous) ,  or  of  carpels  coalescent  into  one  body  (Syncar- 

pous),  or  of  all  grades  between  the  two.     Apocarpous  pistils  are 
simple  ;  a  syncarpous  pistil  is  compound. 

484.  In  both,  the  essential  parts  are  the  ovary  and  the  stigma. 
The  style  may  be  conspicuous  and  widely  separate  these  two, 
as  in  Fig.  536-538  ;  or  hardly  any,  as  in  Fig.  532-535  ;  or  none 
at  all,  as  in  Fig.  530,  531,  533. 

485.  Placenta.     This  name  *  is  applied  to  any  surface  in  the 
interior  of  the  ovary  on  which  ovules  are  borne.     It  has  been 
stated  (579)  that  these  are  usually  borne  upon  the  margins  of 

1  Taken  from  a  remote  analogy  with  the  placenta  of  the  higher  animals. 
The  name  appears  to  have  been  introduced  into  botany  by  Adanson.  It  has 
been  termed  Trophospermum  or  Spermopkorum  by  some  of  the  early  modern 
botanists. 


262 


THE   FLO  WEB. 


the  carpellary  leaf,  or  upon  some  portion  of  what  answers  to 
them.  When  the  ovules  are  numerous,  and  some- 
times when  they  are  few,  the  combined  leaf-edges 
enlarge  to  form  a  kind  of  receptacle  for  their  attach- 
ment or  support :  this  is  the  Placenta.  In  Fig.  530, 
the  placenta  is  well  dev.eloped,  and  also  in  such 
syncarpous  ovaries  as  are  illustrated  in  Fig.  536, 
537,  544,  and  545.  In  very  many  others  (such  as 
Fig.  528,  531,  533),  there  is  no  particular  enlarge- 
ment of  the  leaf-margins  visible,  and  no  particular 
ground  for  the  use  of  this  special  term.  Still  it 
is  commonly  used,  as  occasion  serves,  even  for  the 
mere  line  or  spot  on  which  ovules  are  borne,  as 
well  as  for  a  more  prominent  development  to  which 
the  name  was  originally  applied. 
486.  Simple  or  Apocarpous  Pistils  may  be  solitary,  several,  or 
numerous.  When  indefinitely  numerous,  they  are  seldom  in  one 
circle,  but  are  capitate  or  spicate  upon  a  proportionately  enlarged 
or  prolonged  receptacle,  as  in  Anemone,  Ranunculus,  and  most 
strikingly  in  Myosurus ;  when  reduced  to  a  single  one,  as  in 
Actaea,  Podophyllum,1  Barberry,  and  Plum  or  Cherry,  the  car- 
pel mostly  appears  as  if  it  were  an  actual  termination  of  the 
floral  axis.  But  even  then  the  pistil 
is  hardly  ever  quite  symmetrical  in 
shape  :  the  ovary  is  somewhat  gib- 
DOUS  or  unequal-sided  (as  in  Fig. 

312'315'316'528»531-533)>andthe 
stigma  more  or  less  oblique  or  even 

wholly  lateral.  The  continuation  of 
the  latter  down  the  whole  length  of 
the  ventral  side  of  the  style  (as  in  Fig.  528,  and  also  Fig.  549) 
is  not  uncommon.  In  Schizandra  (Fig.  531)  it  is  continued 
downward  on  the  ventral  edge  of  the  ovary  as  far  as  to  its 
middle.2 


1  Abnormal  specimens  of  Podophyllum  peltatum  are  occasionally  found 
having  a  gynoecium  of  from  two  to  six  separate  carpels. 

2  Pleurogyne,  a  Gentianaceous  genus  so  named  on  this  account,  has  no 
style  nor  apical  stigma  whatever,  but  has  a  long  stigma  extending  down  the 
outside  of  each  ovuliferous  suture  of  its  dicarpellary  ovary  for  most  of  its 
length. 

FIG.  530.    Single  simple  pistil  of  Podophyllum,  cut  across  to  show  the  placenta,  &c. 

FIG.  531.  Vertical  section  of  a  pistil  of  Schizandra  coccinea;  a  side  view  showing 
the  stigma  decurrent  down  to  the  middle  of  the  ovary.  632.  Pistil  of  Hydrastis ;  ventral 
view.  533.  Pistil  of  Actaea  rubra,  cut  across,  so  as  to  show  the  interior  of  the  ovary; 
ventral  view. 


GYNCECIUM  IN  ANGIOSPERMS.  263 

487.  As  the  placenta  of  a  simple  pistil  belongs  to  the  two 
united  margins  of  the  carpellary  leaf,  there  is  naturally  a  double 
row  of  ovules,  one  to  each  margin.     If  the  leaf- 
margins  which  are  turned  inward  in  the  ovary  be- 
low to  bear  the  ovules  are  turned  outward  above  to 

receive  the  pollen  (see  Fig.  531),  then  the  typical 
stigma  should  also  be  double  or  bilamellar.  So  it 
is  seen  to  be  in  such  carpels  as  those  of  Fig.  528, 
531-533,  and  indeed  in  very  many  stigmas  of  this 
class.  Such  division,  or  even  a  greater  bifurcation 
of  a  monocarpellary  stigma  into  two  lobes  or  half- 
stigmas,  is  not  anomalous. 

488.  The  ovary  of  a  simple  pistil  should  be 
unilocular,  that  is,  should  have  a  single   cavity 
or  cell  (loculus),  although,  as  will  soon  be  seen, 
the  converse  does  not  hold  true.     Yet  this  cell  in 
certain  instances  becomes  bilocellate,  being  divided 
by  a  growth  or  intrusion  from  the  back  into  two 
locelli.     This  occurs  more  or  less  in  the  larger 
number  of   species  of   the    Leguminous   genus 
Astragalus,  and  the  mode  is  shown  in  Fig.  534. 

489.  Compound  or  Syncarpous  Pistil.1     This  consists  of  two, 
three,  or  a  greater  number  of  carpels  coalescent  into  one  body. 
A  true  compound  pistil  represents  a  whorl  (in  the  simplest  case 
a  pair)  of  carpels  united  into  one  body,  at  least  as  to  the  ovar}r. 

490.  The  coalescence  of  a  capitate  or  spicate  mass  of  carpels 
or  simple  pistils  of  the  same  flower,  imbricately  heaped  on  the 
torus,   as  in  Magnolia    (Fig.   648)   and   Liriodendron,   cannot 
properly  be   said   to   form   a   compound  pistil.     This  heap  of 
pistils  may  be  called  a  SOREMA. 

491.  Morphologically,  a  compound  pistil,  as  to  the  ovary,  may 
be  a  pair  or  a  circle  of  closed  carpels  or  simple  pistils  brought 
into  contact,  and  the  contiguous  parts  united  :  this  is  illustrated 
in  Fig.  535-538.     Or  it  may  be  formed  of  a  whorl  of  open  car- 
pellary leaves,  joined  each  to  each  by  the  contiguous  margins, 

1  The  terms  apocarpous  and  syncarpous  for  pistils,  the  first  of  separate,  the 
second  of  combined  carpels,  were  introduced  by  Lindley.  They  have  little 
advantage  over  the  terms  simple  and  compound.  Moreover,  the  word 
syncarp  or  syncarpium  had  been  appropriated  to  a  sort  of  fruit  of  the  class 
now  called  multiple,  formed  by  the  coalescence  of  several  flowers,  and  also 
to  that  of  a  heap,  head,  or  spike  of  carpels  more  or  less  cohering  at  matu- 
rity, as  in  a  blackberry,  or  confluent  in  the  flower,  as  in  Magnolia. 

FIG.  534.  Ovary  or  forming  legume  of  Astragalus  Canadensis,  transversely  divided, 
to  show  the  false  partition  which,  intruded  from  the  back,  divides  the  simple  cell  into 
two  half-cells  or  locelli. 


264 


THE   FLOWER. 


in  the  manner  of  Fig.  542-545.     Between  these  two  there  is 
every  gradation.     The  first  forms  a  compound  ovary, 

492.  With  two  or  more  Cells  and  Axile  Placenta.  For  it  is 
evident  that,  if  the  contiguous  parts  of  a  whorl  of  two  or  more 
closed  carpels  cohere,  the  resulting  compound  ovary  should  have 
as  many  cells  as  there  are  carpels  in  its  composition,  and  that 
the  placentae  (one  in  the  inner  angle  of  each  carpel)  will  all  be 


brought  together  in  the  axis  of  the  compound  pistil.  And  the 
partitions,  termed  DISSEPIMENTS,  which  divide  the  compound 
ovary  into  cells,  manifestly  consist  of  the  united  contiguous  por- 
tions of  the  walls  of  the  carpels.  These  necessarily  are  composed 
of  two  layers,  one  belonging  to  each  carpel ;  and  in  fruit  they 
often  split  into  the  two  layers.  True  dissepiments  and  the  true 
cells  must  accordingly  be  equal  in  number  to  the  carpels  of 
which  the  compound  pistil  is  composed.  That  is,  the  ovary,  or 
the  resulting  fruit,  is  Ulocular  or  2-celled,  trilocular  or  3-celled, 
quadrilocular  or  4-celled,  and  so  on,  according  to  the  number  of 
dissepiments  or  cells. 

493.  There  may  also  be  false  dissepiments,  mostly  of  the  same 
character  as  that  which  in  Fig.  534  divides  the  cell  of  a  single 
carpel.  Such  are  found  in  Flax  (Fig.  539-541) ,  in  Amelanchier 
or  Service-berry,  in  Huckleberry  (Gaylussacia) ,  and  in  most  of 


FIG.  635.  Pistil  of  a  Saxifrage  composed  of  two  carpels  or  simple  pistils  united 
below,  but  distinct  above ;  cut  across  both  above  and  below. 

FIG.  636.  Pistil  of  common  St.  Johnswort,  of  three  united  ovaries;  their  styles 
distinct. 

FIG.  637.  The  same  of  another  species  of  St.  Johnswort  (Hypericum  prolificum), 
the  styles  also  united  into  one,  which,  however,  may  split  apart  in  the  fruit. 

FIG.  638.  Pistil  of  Tradescantia  or  Spiderwort,  even  the  three  stigmas  united  into 
one.  The  ovary  in  all  cut  across  to  show  the  internal  structure. 


GYNCECIUM  IN  ANGIOSPERMS. 


265 


the  American  species  of  Vaccinium.  In  all  these,  the  false  par- 
tition is  a  growth  from  the  middle  of  the  back  of  each  carpel, 
which  divides  its  cell  more  or  less  completely  into  two. 


494.  On  the  other  hand,  even  the  true  dissepiments  which 
belong  to  such  a  compound  ovary  may  be  abortive  or  evanescent, 
the  placentae  remaining  in  the  axis  combined  into  a  column. 
(499.)     The  second  modification  of  the  compound  pistil  (491) 
normally  has  an  ovary, 

495.  With   one   Cell   and   Parietal   Placentae.      That    is,   the 
placentae  are  borne  (as  the  term  denotes)  on  the  wall  or  parietes 

of  the  ovary,  as  in  the  Poppy,  Violet,  Sundew, 
Cistus  or  Helianthemum  (Fig.  543),  Cleome,  Gen- 
tian, and  in  all  or  most  of  the  orders  from  which 
these  examples  are  cited.  The  diagram  Fig.  542 
illustrates  the  morphological  conception  of  a  com- 
pound pistil  of  this  kind.  Not  that  it  is  ever  sup- 
posed to  be  formed  by  the  actual  combination  of  once 


separate  leaves,  any  more  than  a  gamophyllous  calyx  or  corolla 
is  actually  so  produced.     The  conception  in  all  such  cases  is  that 

FIG.  639.  Transverse  diagrammatic  section  of  a  flower  of  the  common  Flax,  show- 
ing the  ovary  with  false  partitions  extending  one  from  the  back  of  each  cell.  540.  Sec- 
tion of  a  mature  fruit  and  seeds  of  the  same,  the  false  partitions  now  complete,  divid- 
ing the  five  cells  into  ten.  each  one-seeded.  541.  Same  of  a  wild  Flax  (Linum  perenne), 
in  which  the  false  partitions  remain  incomplete. 

FIG.  542.  Plan  of  a  one-celled  ovary  with  three  parietal  placentae,  cut  across  be- 
low ;  the  upper  part  showing  the  top  of  the  three  leaves  it  is  theoretically  composed  of, 
approaching,  but  not  united. 

FIG.  543.  Ovary  of  Helianthemum  Canadense,  cut  across,  showing  the  ovules  on 
three  parietal  placentae. 

FIG.  544.  Transverse  section  of  the  ovary  of  Hypericum  erraveolens ;  the  three  large 
placentse  meeting  in  the  centre,  but  not  cohering.  546.  Similar  section  of  a  ripe  capsule 
of  the  same;  the  placentae  now  evidently  parietal 


266  THE   FLOWER. 

of  a  congenital  development  of  organs  in  union  which,  in  the 
development  of  a  vegetative  shoot,  would  be  leaves.  This  case 
is  represented  by  the  combination  of  open  carpellary  leaves,  as 
the  preceding  one  is  by  that  of  closed  ones.  As  the  edges 
of  the  leaves  must  needs  be  turned  in,  to  bear  the  ovules, 
a  compound  ovary  with  parietal  placentation  may  be  likened 
to  the  unopened  calyx  of  a  Clematis,  as  shown  in  Fig.  256, 
257.  Every  gradation  is  found  between  axile  and  parietal 
placentation.  Sometimes  the  placentae  are  strictly  on  the  pari- 
etes  or  wall  (Fig.  543,  547)  ;  sometimes  borne  inwards  on 
incomplete  dissepiments  (Fig.  548)  ;  and  sometimes  they  are 
brought  firmly  together  in  the  axis,  as  in  Fig.  544,  though  sepa- 
rable, and  indeed  separated  in  the  fruiting  stage. 

496.  A  compound  ovary  with  parietal  placentas  is  necessarily 
one-celled  (unilocular)  ;  except  it  be  divided  by  an  anomalous 
partition,  such  as  is  found  in  Cruciferae  (Fig.  395)  and  in  many 
Bignoniacese. 

497.  Normal  placentae  are  necessarily  double  :  when  parietal, 
the  two  halves  belong  to  different  leaves  ;  when  axile,  to  the  same 
leaf.     These  two  halves  may  diverge  or  be  widely  separated, 


sometimes  even  at  their  origin,  as  in  Aphyllon  and  some  other 
Orobanchaceae,  in  which  a  dicarpellary  ovary  has  four  almost 
equidistant  placentas ;  or  in  such  cases  the  placentae  may  be 
regarded  as  intra-marginal  instead  of  marginal. 

498.  The  placentae  of  a  two-several-celled  ovary,  such  as  in 
Fig.  536,  537,   &c.,  may  be  described  in  the  plural  number, 
being  one  in  each  carpel ;  or  when  consolidated  into  a  central 
column,  and  well  covered  with  ovules,  they  ma}'  be  said  to  form 
one  (compound)  placenta.     Then  when  the  dissepiments  early 
disappear,  or  are  abortive  from  the  first,  the  result  is  a  compound 
ovary  of  this  class, 

499.  With  one  Cell   and  Free   Central  Placenta.     In  Caryo- 
phj-llaceae  (Fig.   549,   550)    and   Portulacaceae,  this   evidently 
results  from  the  obliteration  of  the  dissepiments  (as  many  as 
there  are  styles  or  stigmas) ,  vestiges  of  which  may  be  sometimes 

FIG.  546.  Diagram  (ground-plan)  to  illustrate  free  central  placentation  produced 
by  abortion  of  dissepiments.  547.  Same  of  strict  parietal  placentation.  548.  Same  with 
the  placentae  carried  inward  on  imperfect  dissepiments.  , 


GYNCECIUM  IN   ANGIOSPERMS.  267 

detected,  while  certain  plants  of  the  same  families,  of  otherwise 
identical  structure,  retain  the  dissepiments  even  in  the  fruit. 

500.  But  a  similar  condition  may  equally  arise 
from  a  modification  of  parietal  place ntation,  namely, 
with  the  margins  of  the  leaves  ovuliferous  only  at 
bottom,  and  the  placentae  there  conspicuously  devel- 
oped and  completely  united.     The  basal  placenta- 
tion  of  Dionaea  is  unavoidably  so   explained,  its 
nearest  relative,  Drosera  (Fig.  553) ,  having  parietal 
placentas.     And  this  leads  to  a  probable  explanation 
of  the  case  in  Primulacese,  where  a  large  free  central 
placenta  fills  the  centre  of  the  cell,  and  no  trace  of 
dissepiments  can  be  detected.1 

501.  The  idea  maintained  in  former  editions  is 
still  adhered  to ;  namely,  that  placentae  belong  to 
carpels  and  not  to  the  cauline  axis,  in  other  words, 

that  ovules  are  productions  of  and  borne  upon  leaves,  usually 
upon  their  margins,  not  very  rarely  upon  other  portions  of  their 
upper  surface,  rarely  over  the  whole  of  it.2 

502.  Ovules  cover  the  whole  internal  face  of  the  carpels  in 
Butomus  and  its  relatives,  also  of  the  "Water-Lilies  (both  Nym 
phaea  and  Nupliar,  Fig.  551)  excepting  the  inner  angle,  to  whicl* 
they  are  usually  restricted  in  other  plants.     And  in  the  allied 
Brasenia  and  Cabomba,  where  the  ovules  are  reduced  to  two  or 
three,  one  or  more  of  them  is  on  the  midrib,  but  none  on  the 

1  The  placenta  in  this  and  like  cases  is  rather  to  be  regarded  as  an  out- 
growth from  the  base  of  the  carpellary  leaves,  combined  over  the  floral 
axis.     Upon  this  interpretation,  a  central  portion  of  the  column  may  be 
(and  sometimes  must  be)  of  axile  nature,  yet  the  ovules  be  borne  upon 
foliar  parts.     See  Van  Tieghem,  in  Ann.  Sci.  Nat.  ser.  5,  xii.  329  (1869) ; 
Celakowsky,  Vergleichende  Darstellung  der  Placenten,  &c.  (1876) ;  Warming, 
in  Ann.  Sci.  Nat.  ser.  6,  v.  192. 

2  This  view  was  first  maintained  as  a  general  theory,  and  on  critical 
grounds,  by  Brown,  in  Plantae  Javanicae  Rariores,  107-112.     Schleiden,  End- 
licher,  and  others  took  the  opposite  view,  i.  e.,  that  ovules  are  productions  of 
the  axis,  even  in  parietal  placentation,  —  an  exceedingly  far-fetched  suppo- 
sition.    In  later  days,  the  commoner  view  has  regarded  ovules  as  of  both 
origins,  as  productions  of  the  carpels  in  parietal,  of  the  axis  in  at  least 
some  free   central  or  basilar  placentation.     But  at  present  the  theory  of 
foliar  origin  without  exception,  revindicated  by  Van  Tieghem,  and  espe- 
cially by  Celakowsky  and  Warming,  again  prevails.     For  the  bibliography 
and  an  abstract  of  the  various  views,  see  Eichler,  Bliithendiagramme,  espe- 
cially the  note  in  the  preface  to  the  second  part  (where  he  gives  his  entire 
adhesion  to  this  conclusion)  ;  also  Warming's  memoir,  De  1'Ovule,  in  Ann. 
Sci.  Nat.  ser.  6,  v.  1877-78. 

FIG.  549.    Vertical  section  through  the  compound  tricarpellary  ovary  of  Spergularia 
rubra,  showing  the  free  central  placenta.    550.  Transverse  section  of  the  same. 


THE   FLOWER. 


margins  of  the  carpellary  leaf.  In  many  species  of  Gentian,  as 
also  in  Obolaria  and  Bartonia,  of  the  same 
family,  the  whole  internal  face  of  a  dicar- 
pellary  ovary  is  thickly  ovuliferous. 

503.  Perhaps  the  parietal  placentte  in 
Parnassia  (Fig.  552)  are  borne  on  the 
midribs  of  the  carpels,  for  they  are  directly 
under  the  stigmas,  instead  of  alternate 
with  them,  as  they  normally  should  be.  The  same  thing  occurs 
in  Poppies  and  many  other  Papaveracese,  also  in  some  Cruciferae  ; 
and  in  some  of  the  cases  each  stigma 
is  more  or  less  two-lobed.  This  sug- 
gests the  explanation,1  here  probably 
the  true  one,  which  supposes  that  the 
placentae  are  borne  on  the  leaf-margins 
in  the  normal  way,  but  that  each 
stigma  is  two-parted  (as  if  the  carpel- 
lary leaf  were  deeply  notched  at  the 
apex,  and  so  its  two  stigmatic  leaf- 
margins  separate,  as  Drosera  illus- 
trates, Fig.  553),  and  that  the  two 
half-stigmas  of  adjacent  carpels  have  553 

coalesced  into  one  body,  which  would 
of  course  stand  over  the  parietal  placentae  beneath.    Each  stigma 
in  such  a  case,  as  well  as  each  parietal  placenta,  would  consist 
of  the  united  margins  of  two  adjacent  carpels. 


§  2.    IN  GYMNOSPERMS. 

504.  GYMNOSPERMOUS  (that  is,  naked-seeded)  plants  are  so 
named  because  the  ovules,  or  bodies  which  are  to  become  seeds, 
are  fertilized  by  direct  application  of  the  pollen,  which  reaches 
and  acts  upon  the  nucleus  of  the  ovule  itself,  not  through  the 
mediation  of  stigma  and  style.  In  the  structure  of  their  flowers, 
these  plants  are  of  a  low  or  simplified  type,  in  some  respects  not 
obviously  homologous  with  the  Angiospenns  which  now  consti- 
tute the  immense  majority  of  phaenogamous  plants.  But,  up  to 
a  comparatively  late  geological  period,  Gymnosperms  appear  to 
have  been  the  only  flower-bearing  plants.  They  are  represented 

1  Given  by  Brown,  in  the  Plantse  Javanicae  Rariores,  above  referred  to. 

FIG.  551.  Transverse  section  of  an  ovary  of  Nymphsea  odorata,  the  carpels  ovulifer- 
ous over  the  whole  Interior  surface. 

FIG.  552.    Pistil  of  Parnassia,  with  ovary  transversely  divided. 

FIG.  663.    Piatll  of  Drosera  filiformis,  with  ovary  transversely  divided. 


GYNCECITJM  OF   GYMNOSPERMS.  269 

in  the  extant  vegetable  kingdom  by  three  (or  four)  groups  or 
orders,  two  of  them  small,  and  one  comparatively  ample  and  of 
wide  distribution ;  and  all  are  so  strikingly  different  from  each 
other  that  they  cannot  be  illustrated  by  a  common  description. 
The  largest  order,  Coniferae,  is  familiar,  and  contains  a  good 
share  of  the  most  important  forest  trees  of  temperate  climates. 
The  smallest,  Gnetaceae,  chiefly  tropical  or  of  warm  regions, 
lies  between  Gymnosperms  and  common  Dicotyledons.  The 
third,  Cycadacese,  is  most  remote  from  them,  and  as  much  so 
from  Monocotyledons,  except  that  it  imitates  Palms,  as  it 
also  does  the  Tree-Ferns,  in  habit,  both  as  to  stem  and  foli- 
age. The  particular  morphology  of  Gymnosperms  would  re- 
quire for  its  illustration  copious  details  and  the  history  of  various 
conflicting  hypotheses.  It  must  be  relegated  to  the  special 
morphology  of  the  natural  orders,  premising,  however,  a  brief 
Sketch  of  the  general  floral  structure.1 

505.  In  Gnetaceae,  Gymnosperms  and  Angiosperms  almost 
tome  together.  The  flowers  have  a  perianth  (diphyllous  or 
tetraphyllous)  ;  the  stamens  have  a  distinct  filament  and  anther ; 
and  the  gyncecium  is  a  sac  (presumably  of  two  carpophylls) 
open  at  the  top  and  filled  at  bottom  by  a  single  ovule  of  the 
simplest  kind,  «'.  e.  consisting  of  a  nucleus  destitute  of  coats. 
This  pistillary  body  is  attenuated  and  prolonged  above  the  ovule 
into  a  style-shaped  tube,  with  open  and  commonly  two-cleft 
orifice.  In  the  almost  hermaphrodite  sterile  flower  of  Welwitschia, 
this  takes  the  form  of  a  much  dilated  stigma,  which  is  even  beset 
with  seeming  stigmatic  papillae.  If  only  the  pollen  were  here 
to  grow  forth  into  pollen-tubes  (with  or  without  a  closing  of  the 
tube) ,  angiospermy  would  be  attained.  But,  in  fact,  the  pollen- 
grains  bodily  reach  the  ovule  itself  through  the  tube,  fertilizing 
it  directly.'2  This  interesting  group  of  plants  consists  of  the 

1  References  to  the  literature  of  gymnospermy  and  to  the  steps  of  the 
prolonged  controversy  over  it,  also  the  points  of  morphology  still  in  part 
unsettled,  need  not  here  be  given.     The  history  and  the  idea  of  gymnospermy 
began  with  Robert  Brown's  paper  on  Kingia,  "  with  Observations  ....  on 
the  Female  Flower  of   Cycadeae  and  Coniferae,"  read  before  the  Linnean 
Society  in  the  year  1825,  and  published  in  King's  Voyage  in  1827  ;  and  the 
bibliography  down  to  a  recent  date  is  given  by  Eichler  in  Flora  Brasiliensis, 
Gymnospermia,  iv.  435,  and  in  Bliithendiagramme,  i.  55-69 ;  also  ii.  preface  x. 
See  also  Alph.  DeCandolle,  Prodr.  xvi.2  345,  524.    In  this  volume,  the  late 
Prof.  Parlatore  adhered  to  the  ancient  ideas  in  his  monograph  of  the  Coniferae. 

2  The  view  here  implicitly  adopted  is  that  of  Beccari,  founded  on  the  study 
of  Gnetum,  and  published  in  Nuovo  Giornale  Botanico  Italiano,  ix.  1877. 
It  was  before  nearly  or  quite  reached  in  successive  steps,  by  J.  D.  Hooker, 
in  his  classical  memoir  on  Welwitschia,  in  Trans.  Linn.  Soc.  xxiv.;  Stras- 
burger,  Die   Coniferen  und  die   Gnetaceen,   1872;  and  W-  R.  McNab.  in 
Trans.  Lin»  Sf>c.  xxviii.  1872. 


270 


THE   FLOWER. 


genus  Gnetum,  shrubs  or  trees,  with  nearly  the  aspect  of 
Angiospenns,  having  broad  and  pin nately- veined  leaves ;  Wel- 
witschia  of  tropical  W.  Africa,  remarkable  for  its  persistent 
cotyledons  which  form  the  only  foliage  of  a  woody  and  long- 
enduring  plant,  and  for  its  stem  or  trunk  which  broadens  with- 
out lengthening,  except  in  its  flower-stalks  ;  also  Ephedra,  of 
much  branched  shrubs,  mainly  of  warm-temperate  regions,  leafless 
or  nearly  so,  one  species  of  which  inhabits  Europe  and  two  the 
southern  borders  of  the  United  States. 

506.  The  flowers  in  all  Gymnosperms   are   diclinous,  either 
dioecious  or  monoecious  ;  except  that  those  of  the  strange  Gneta- 
ceous  genus  Welwitschia  are  structurally  polygamous,  the  male 
flowers  having  a  well-formed  but  sterile  gynoecium. 

507.  In  Coniferae,  the  largest  and  most  important  type,  are 
embraced  all  the  familiar  Gymnosperms  of  temperate  regions, 

Pines,  Firs,  Cedars,  Cypresses,  which 
bear  their  flowers  in  catkin-like  clusters 
and  their  fruit  in  cones,  and  also  the 
Yews  and  allied  trees  which  do  not 
produce  cones.  Perianth  being  want- 
ing and  the  sexes  wholly  separate,  the  floral  type 
is  so  degraded  that  it  becomes  doubtful  whether 
each  cluster  of  anthers,  or  of  ovuliferous  scales 
or  ovules,  constitutes  a  blossom  or  an  inflores- 
cence. Certain  botanists  look  upon  a  whole 
catkin,  and  others  upon  a  male  catkin  only,  of 
a  Pine  or  Fir  as  forming  one  flower.  It  is  here 
assumed  that  each  stamen 
of  the  one  and  each  ovu- 
liferous scale  of  the  other 
answers  to  a  flower  of  the 
simplest  sort.1  The  anthers 
555  are  extrorse,  the  cells  or 

pollen-sacs  belonging  to  the  outer  or  lower  side  of  a  scale  or  a 

1  It  will  be  seen  that,  for  the  female  flowers,  this  follows  of  course  from 
generally  accepted  view ;  and,  where  this  is  conceded,  analogy  may  extend  it 
to  the  male  catkins  also:  yet  in  such  cases,  where  all  the  phylla  of  an  indefinite 
simple  axis  are  stamens,  spirally  arranged  on  it,  the  difference  between 
inflorescence  and  male  flower  completely  vanishes. 

FIG.  554.  Female  flower  of  a  Yew,  an  ovule  surrounded  by  its  bracts.  555.  Longi- 
tudinal and  more  enlarged  section  of  a  female  flower  of  Yew  and  of  the  tipper  part  of 
the  shoot  it  terminates :  the  thick  coat  of  the  ovule  open  at  the  top,  the  nucleus  within, 
and  the  beginning  of  the  disk  outside  of  the  coat,  are  seen  in  section.  After  Stras- 
burger. 

FIG.  556.  Young  fruit  (berry-like  cup  surrounding  the  seed)  of  Yew.  557.  Longi- 
tudinal section  of  a  mature  fruit  of  the  same.  After  Decaisne. 


GYNCECIUM  OF   GYMNOSPERMS. 


271 


connective  :  sometimes  these  sacs  or  cells  are  two,  and  the  organ 
evidently,  homologous  with  an  ordinary  stamen :  often  they  are 
more  numerous  (from 
three  to  twenty)  and 
variously  disposed. 

508.  The  Yew  Fam- 
ily (Taxinese)  is  next  to 
Gnetaceae  in  structure. 
It  is  generally  ranked  as 
a  suborder  of  Coniferse, 
but  it  may  claim  to  be  a  distinct  order.  The  gynrecium  is  a 
naked  ovule,  terminating  a  stem,1  and  surrounded  by  several 
bracts.  After  fertilization,  an  outgrowth  of  the 
receptacle  (or  a  kind  of  disk,  394)  makes  its 
appearance  as  a  ring  girding  its  base :  this 
grows  in  height  and  thickness,  and  becomes  a 
soft-fleshy  cup,  imitating  a  hollow  berry,  in 
the  bottom  of  which  the  stony-coated  seed 
nestles.  (Fig.  554-557.)  Very  similar  is  the 
gynoecium  of  Torreya,  except  that  the  cup- 
shaped  disk  develops  almost  simultaneously 
with  the  ovules,  and  as  it  grows  becomes  adnate 
to  the  large  seed  in  the  form  of  a 
fleshy  coating.  In  the  Gingko,  two 
or  more  similar  ovules  are  nakedly 
developed  on  a  naked  peduncle,  un- 
accompanied even  by  a  bract  (Fig. 
558) ,  and  one  or  more  of  these  ripens 
into  the  berry-like  seed,  Fig.  559. 
In  Podocarpus  there  are  some  sub- 
tending bracts,  and  the  naked  ovule 

1  It  does  not  therefore  follow  that  the  ovule  is  a  part  of  the  axis,  or  is 
terminal  in  the  sense  of  being  its  direct  continuation.  In  this  regard  it  may 
be  only  what  the  pistil  of  a  Cherry  is,  which  to  all  appearance  is  equally  a 
terminal  production,  but  is  really  the  representative  of  the  last  leaf  of  the 
axis.  If  so,  that  leaf  is  here  suppressed  to  the  utmost,  and  replaced  by 
what  is  ordinarily  its  outgrowth,  the  ovular  nucleus  and  its  coat.  The 
structure  of  Podocarpus  favors  this  interpretation. 

FIG.  558.  Female  flowers  of  Gingko  biloba  or  Salisburia  adiantifolia.  558o.  Portion 
of  the  same  enlarged.  After  Strasburger.  559.  A  drupaceous  seed  of  the  same,  in 
vertical  section,  exhibiting  the  mature  disk  which  forms  the  flesh,  the  crustaceous  seed- 
coat,  within  which  is  the  kernel  of  the  seed ;  at  the  base  on  one  side  a  sterile  ovule  is 
seen.  After  Decaisne. 

FIG.  560.  Female  flower  of  Podocarpus  (an  ovule  inverted  on  a  column  or  elevated 
support),  subtended  by  bracts.  After  Eichler. 

FIG.  561.  Magnified  vertical  section  of  &  similar  flower  of  Podocarpus.  After 
Strasburger. 


272 


THE   FLOWER. 


is  inverted  on  a  more  or  less  lengthened  and  stout  support,  which 
is  conceived  to  represent  the  carpel.  (Fig.  560,  561.) 

509.  In  the  true  Coniferae,  to  which  Pines,  Cypresses,  and  all 
such  cone-fruiting  trees  belong,  the  ovules  are  borne  on  or  in 
the  axils  of  scales  which  are  imbricated  on  a  simple  axis,  in  a 
spicate  or  capitate  manner  ;  and  the  male  flowers,  each  a  single 
stamen,  are  also  similarly  spicate  or  capitate.     Both  are  com- 
monly termed  aments  or  catkins ;  and  the  female 
ones  properly  so,  according  to  the  present  view ; 
but  the  only  scales  of  the  male  catkins  are  parts 
of  the  anther,  being  a  dilated  tip  of  the  connective 
in  Pines,  and  a  scale  bearing  anther-cells  or  pollen- 

562  sacs  on  its  back  in  Cypress. 

510.  In  the  Pine  tribe  the  flowering  female  catkin  consists  of 
bracts,  spirally  imbricated  on  the  cauline  axis :  in  the  axil  of 

each  bract  or  sterile  scale  is  developed  a  scale 
which  bears  two  ovules,  and  is  therefore  regarded 
as  of  carpellary  nature.  These  ovules  are  pro- 
duced on  the  lower  part  of  the  upper  face  of  this 
carpellary  scale,  and  are  wholly  adherent  to  it 
quite  to  the  orifice,  which  is  directed  downward. 
(Fig.  562,  563.)  The  ovuliferous  scale  in 
becoming  fructiferous  usually  much  and  soon  out- 
grows the  bract,  which  is  concealed  in  the  Pine- 
cone  (or  sometimes  obliterated) ;  but  it  remains 
conspicuous  in  sundry  Fir-cones.  After  fertil- 
ization, the  scales,  successively  covering  each 
other  in  close  imbrication,  protect  the  growing 
seeds  as  effectually  as  would  a  closed  ovary. 
Sooner  or  later  after  ripening  the  scales  diverge,  and  the  seeds 
peel  off  the  face  of  the  scale  with  a  wing  attached,  and  fall  or 
are  dispersed  by  the  wind.1 


1  Among  those  who  admit  as  well  as  those  who  reject  gymnosperrc^ , 
there  has  been  much  controversy  over  the  morphology  of  the  parts.  With 
the  former,  the  discussion  turns  on  the  character  of  the  ovuliferous  scale. 
As  to  this,  the  hypothesis  originally  proposed  by  Mohl,  and  adopted  by 
Braun,  is  now  said  to  be  satisfactorily  demonstrated  by  Stenzel,  in  Nov.  Act. 
Nat.  Cur.  xxxviii.  1876.  See  note  by  Engelmann  in  Amer.  Jour.  Sci.  Dec. 
1876,  and  also  the  preface  to  the  second  part  of  Eichler's  Bliithendiagramme, 

FIG.  562.  View  of  the  upper  face  of  a  carpellary  scale  of  a  Larch,  showing  the  pair 
of  adnate  ovules. 

FIG.  563.  Similar  view  of  a  carpellary  scale  of  a  Larch,  and  of  a  bract  behind  it. 
564.  Ground  plan  of  the  same  in  diagram,  reversed;  the  upper  figure  denoting  the  axis 
of  the  cone,  the  lower  the  bract,  the  middle  one  the  carpellary  scale  and  the  two  ovule* 
borne  on  its  face.  After  Eichler. 


GYNCECIUM   OF  GYMKOSPERMS. 


273 


511.  In  the  Araucaria  tribe  the  ovuliferous  or  carpel-scale  is 
throughout  smaller  than  the  bract,  and  is  completely  adnate  to 
it,  or  with  only  the  tip  free  ;  that  of  Araucaria  (Fig.  565)  bears 
only  one  ovule,  high  on  the  carpel,  the  orifice  downward  as  in 
the  Pine  tribe.     In  Taxodium,  Sequoia,  and  the  like,  the  cone- 
scale  is  equally  inferred  to  be  composed  of  bract  and  carpel-scale 
united  ;  and  indications  of  this  composition  are  to  be  observed. 
The  ovules  (from  two  to  several)  are  at  the  base  of  the  scale, 
erect  and  free.     The  cone-scales  are  alternate  and  spiral  on  the 
axis,  but  indistinctly  so   in   Taxodium,   the   Bald   Cypress  or 
so-called  Cypress  of  the  Southern  United  States. 

512.  In  the  true  Cypress  tribe  (Cupressineae)  the  cone-scales, 
which  are  never  numerous,  are  opposite  or  verticillate,  i.  e.  like 
the  foliage- leaves,  in  whorls  of  twos,  threes,  or 
sometimes  fours  ;  and  the  ovules  are  from  two  to 


1878,  where  it  is  fully  adopted.    It  was  suggested  by  certain  rather  common 
monstrosities,  and  by  the  two  combined  leaves  of  Sciadopitys. 

According  to  this  view,  the  ovuliferous  scale  in  the  Pine  tribe  is  com- 
posed of  two  leaves  of  an  arrested  and  transformed  branch  from  the  axil  of 
the  bract,  which  are  in  the  normal  manner  transverse  to  the  subtending 
bract,  are  here  carpellary,  each  bearing  an  ovule  on  the  dorsal  face ;  the  two 
are  coalescent  into  one  by  the  union  of  their  posterior  edges,  and  the  scale 
thus  formed  is  thus  developed  with  dorsal  face  presented  to  the  axis  of  the 
cone,  the  ventral  to  the  bract.  It  is  therefore  a  compound  open  carpel, 
composed  of  two  carpophylls.  This  character  of  being  fructiferous  on  the 
back  or  lower  side  of  the  leaf  occurs  in  no  other  phaenogamous  plants,  but  is 
the  rule  in  Ferns,  from  something  like  which  Coniferae  may  be  supposed  to 
have  been  derived ;  the  ovules  of  the  one  in  this  regard  corresponding  to  the 
sporangia  of  the  other. 

FIG.  565.  Vertical  section  (in  diagram)  of  a  bract,  adnate  carpel-scale,  and  adnate 
ovule  of  Araucaria  imbricata.  After  Eichler. 

FIG.  566.  Branchlet  of  the  American  Arbor-Vitas,  considerably  larger  than  in  na- 
ture, with  a  forming  fertile  cone.  567.  One  of  the  scales  removed  and  more  enlarged, 
the  inside  exposed  to  view,  showing  a  pair  of  naked  erect  ovules  on  its  base. 

FIG.  568.  Fertile  flowers  of  true  Cypress  (Cupressus  seaipervirens),  after  Baillon :  a 
forming  cone,  with  one  scale  cut  away,  to  show  the  cluster  of  ovules  under  it. 

18 


274 


THE  FLOWER. 


several  at  or  on  the  base  of  each  cone-scale,  always  with  orifice 
upward.  Arbor- Vitse  (Fig.  566,  567)  has  a  single  pair  of  ovules 
to  the  scale  ;  Junipers,  sometimes  only  one ;  true  Cypresses  (as 
in  Fig.  568),  often  a  dozen  or  more.  At  flowering  time,  the  cone- 
scales  mostly  appear  as  if  simple  ;  but  in  most  genera  they  soon 
thicken  greatly  within ;  and  they  are  usually  understood  to  be 
composed  of  bract  and  carpel-scale  combined,  the  latter  of  the 
same  constitution  as  that  of  Pines  and  Spruces,  but  perfectly 
consolidated  and  confluent  with  the  bract-scale.1 


513.  In  Cycadacese,  the  type  of  the  flower  of  Angiosperms  is 
almost  or  quite  lost ;  yet  the  organs  may  be  homologized  with 
those  of  Conifera3,  which  these  plants  are  wholly  unlike  in  habit. 


1  This  internal  and  ovuliferous  scale  may  seem  to  be  wholly  hypotheti- 
cal, and  assumed  to  homologize  the  cupressineous  with  the  abietineous 
cone.  Without  it,  we  should  have  to  consider  that,  while  in  Abietineae  the 
ovules  belong  to  leaves  of  a  secondary  axis,  in  Cupressineae  they  are  borne 
on  those  of  a  primary  axis,  or  else  are  axillary  productions  without  carpels. 
But  in  the  Araucaria  tribe  the  internal  scale  is  obvious  ;  and  there  are  suffl- 

FIG.  569-575.  Zamia,  chiefly  Z.  media,  after  Richard.  569.  A  male  plant,  570 
Lower  part  of  a  male  catkin.  571.  A  stamen  removed,  showing  numerous  small  pollen- 
sacs  under  the  peltate  top.  572.  A  female  catkin,  with  a  quarter  section  cut  away. 
573.  A  female  flower  or  carpel,  with  two  enlarging  ovules  or  young  seeds.  574.  Bine 
seed,  with  the  thick  fleshy  coat  cut  away  at  apex.  575.  Longitudinal  section  of  ripe 
iied,  more  enlarged. 


GYNCECIUM   OF   GYMNOSPEKMS. 


2T5 


Their  likeness  to  Palms  and  other  Monocotyledons  is  confined  to 
the  port  of  their  unbranched  trunks  and  their  pinnate  leaves 
with  parallel-veined  or  simple- veined  leaflets  ;  nor  have  they  any 
further  resemblance  to  Ferns,  except  that  in  some  the  leaflets  are 
circinate  in  vernation.  Although  a  tropical  type  (of  small  present 
importance,  compared  with  the  part  which  it  played  in  the 
Devonian  and  Cretaceous  periods) ,  it  has  one  small  representa- 
tive (Zamia  media,  the  Coontie)  at  the  south-eastern  extremity  of 


the  United  States,  and  a  more  striking  one  (Cycas  revoluta,  well 
known  in  cultivation)  in  the  southern  parts  of  Japan. 

514.  Following  the  analogy  of  Coniferse,  each  scale  (whether 
of  the  pollen-bearing  or  the  ovule-  and  seed-bearing  ament)  of 
Zamia  (Fig.  569-575)  is  here  regarded  as  a  flower.  Here  the 
phylla,  or  scales  with  peltate  top  and  stalk-like  base,  are  exter- 


cient  indications  of  similar  composition  in  the  cupressineous  cone-scales  to 
induce  the  adoption  of  it  by  Parlatore,  who  rejected  the  idea  of  gymnospermy ; 
and,  finally,  this  composition  is  nearly  demonstrated  by  VanTieghem  (1868) 
upon  the  anatomical  structure,  and  by  Strassburger  ( 1872)  on  the  development 

FIG.  576-578.  Carpophylla  of  Cycas  revoluta,  much  reduced  In  size.  P76.  One 
bearing  ovules  below  and  leaflets  or  leaf-lobes  towards  the  apex.  577.  A  similar  carpo- 
phyll  with  leaf-lobes  reduced  to  mere  teeth,  and  ovules  in  place  of  the  lower  teeth. 
578.  A  similar  carpophyll  in  mature  fructification,  bearing  the  large  drupaceous  naked 
seeds.  The  last  two  after  Richard. 


276  THE  FLOWER. 

nally  much  alike  in  the  two  sexes,  which  throughout  the  family 
occupy  separate  plants.  The  male  flower  (Fig.  568)  or  stamen, 
if  it  may  be  so  termed,  bears  indefinite  pollen-sacs  on  the  under 
side  of  the  peltate  portion,  sometimes  extending  to  che  upper 
part  of  its  stalk.  The  homologous  female  flower,  or  carpophyll, 
bears  a  suspended  ovule  on  each  side  of  the  stalk  (Fig.  573), 
which  becomes  a  large  fleshy-coated  seed.  In  Cycas  the  male 
ament  is  not  very  dissimilar,  although  on  a  larger  scale.  But 
the  carpophylls  are  evident  leaves,  not  condensed  into  an  ament, 
but  loose  or  spreading,  of  a  character  and  aspect  intermediate 
between  the  lax  bud-scales  which  precede  and  the  pinnate  foliage- 
leaves  which  follow  them  in  development.  Along  the  margin 
of  what  would  be  leaf-blade  they  bear  ovules  in  place  of  leaflets, 
lobes,  or  teeth  (Fig.  576-578)  ;  and  these,  when  fertilized  from 
the  male  flowers,  mature  into  large  and  drupaceous  naked  seeds. 
Even  without  fertilization,  such  seeds  grow  to  their  full  size  on 
the  female  plant  of  the  common  Cycas  (or  falsely  so-called  Sago 
Palm) ,  but  form  no  embryo. 


SECTION  VHI.    THE  OvuLE.1 

515.  Cynics  (302)  are  peculiar  outgrowths  or  productions  of 
carpels  which,  upon  the  formation  of  an  embryo  within,  become 
seeds.     In  the  angiospermous  gynoecium  (476)  they  are   nor- 
mally produced  along  the  margins,  or  some  part  of  the  margins, 
of  the  carpellary  leaf  (478) ,  either  immediately,  or  by  the  in- 
termediation of  a  placenta  (485),  which  is  a  more  or  less  evident 
development  of  the  leaf-margins  for  the  support  of  the  ovules. 
Rarely,  yet  in  a  considerable  number  of  cases  (501,  502),  ovules 
are  developed  from  the  whole  internal  surface  of  the  ovary,  or 
from  various  parts  of  it,  in  no  definite  order,  directly  from  the 
walls,  and  without  the  intervention  of  any  thing  which  can  be 
regarded  as  placenta.     In  Gymnosperms  (504-514)  the  ovules 
are  borne  on  the  face  of  the  carpellary  scale  or  at  its  base ;  or 
on  leaf-margins,  as  in  Cycas ;  or,  when  there  is  no  representa- 
tive of  the  carpel,  on  the  cauline  axis,  seemingly  as  a  direct 
growth  of  it.     (508,  note.) 

516.  As  to  attachment,  ovules  are  either  sessile,  i.  e.  stalk- 
less,  or  on  a  stalk  of  their  own  (Fig.  582,  584),  the  FUNICULUS 
or  PODOSPERM.     As  to  number  they  are  either  solitary,  few,  or 

1  Lat.  Ovtdum,  pi.  Ovtda,  diminutive  of  ovum  (egg),  perhaps  first  used  by 
Adanson. 


OVULES. 


277 


indefinitely  numerous.     They  may  also  be  indefinite  or  variable 
in  number  when  not  particularly  numerous. 

517.  As  to  situation  and  direction  within  the  ovary,  the  terms 
are  somewhat  special.     Ovules  are  erect,  when  they  rise  from 
the  very  bottom  of  the  cell,  as 

in  Fig.  580 ;  ascending,  when 
attached  above  its  bottom  and 
directed  upward,  as  in  Fig.  579  ; 
horizontal,  when  borne  on  one  or 
more  sides  of  the  cell  and  not 
directed  either  upward  or  down- 
ward, as  in  Fig.  314,  315,  530 ; 
pendulous,  when  more  or  less  hanging  or  declining  from  the  side 
of  the  cell ;  suspended,  when  hanging  from  the  apex  of  the  cell, 
as  in  Fig.  581. 

518.  The  body  and  only  essential  part  of  an  ovule  is  its 
NUCLEUS.     This  in  most  cases  is  invested  by  one  or  two  proper 
coats.     The  coats  are  sacs  with  a  narrow  orifice,  the  FORAMEN. 
In  the   seed,  the  closed  vestige  of  this  orifice  is  termed  the 
Micropyle ;  wherefore  this  name  is  sometimes  applied  to  it  in 
the  ovule  likewise.     When  the  ovule  has  two  coats,  the  foramen 
of  the  outer  one  is  called  EXOSTOME,  of  the  inner  ENDOSTOME  ; 
literally  the  outer  and   the   inner 

orifice.    The  coats  themselves  have 

been  named  PRIMINE  and   SECUN- 

DINE,  but  with  an  ambiguity  in  the 

application    which    renders    these 

names    unadvisable :    for  in  their 

formation  the   coats   appear  later 

than  the   nucleus,  the   inner  coat 

earlier  than   the    outer;   and   the 

name    of   primine    has    by    some 

writers  been  applied  to  the  earlier 

formed,  by  others  to  the  external  coat.     The  proper  base  of  the 

ovule,  from  which  the  coats  originate  and  where  these  and  the 

nucleus  are  confluent,  is  the  CHALAZA.     The  attachment  of  the 

ovule  to  its  funiculus  or  support,  which  in  the  seed  becomes  the 


FIG.  579.  Ovary  of  a  Buttercup,  divided  lengthwise,  to  display  its  ascending  ovule. 
680.  Same  of  Buckwheat,  with  an  erect  ovule.  581.  Same  of  Anemone,  with  a  sus- 
pended ovule. 

FIG.  582.  Diagrammatic  section  of  a  typical  or  orthotropous  ovule  (such  as  that  of 
Kg.  582"),  showing  the  outer  coat,  a,  the  inner,  b,  the  nucleus,  c,  the  chalaza,  or  place 
of  junction  of  these  parts,  d.  (The  coats  are  never  so  separated  and  the  nucleus  so  re- 
duced in  size  as  is  represented  in  this  mere  diagram.)  583.  An  ovule  similar  to  the 
preceding,  but  curved,  or  campylotropous.  684.  An  amphitropous  ovule. 


278  THE  FLOWER. 

HILUM,  takes  also  this  latter  name  in  the  ovule.  In  the  simplest 
form  of  ovule  (as  in  Fig.  582,  580),  hilum  and  chalaza  are  one. 
So  also  in  cases  where  the  body  of  the  ovule  incurves,  as  in  Fig. 
583.  But  very  commonly  the  place  of  attachment,  which  becomes 
the  hilum,  is  more  or  less  distant  from  the  chalaza  ;  as  in  Fig.  584 
and  587,  where  the  hilum  is  lateral,  but  the  chalaza  at  the  larger 
end,  the  two  being  connected  by  a  short  ridge  ;  and  in  Fig.  588 
the  two  are  separated  by  the  whole  length  of  the  ovule. 

519.  The  simplest  and  most  rudimentary  ovule  is  that  with- 
out a  coat,  as  in  Mistletoe  and  the  whole  order  Loranthaceae,  and 
in  Santalaceae  and  Gnetaceae.     This  has  been  called  a  naked 
ovule ;   but  long  before  ovules  of  such  simplicity  were  known 
this  term  had  been  appropriated  to  those  of  Gymnosperms,  in 
the  sense  of  destitute  of  ovarial  or  pericarpial  covering,  i.  e.  to 
uncovered  ovule,  not  to  uncovered  nucleus.     The  ovule  consist- 
ing only  of  nucleus  may  be  termed  (after  Alph.  DeCandolle) 
simple,  or  better  achlamydeous.1 

520.  The  tunicated  or  chlamydeous  ovule  is  of  three  principal 
kinds,  with  one  or  two  subordinate  modifications.     These  are 
the  Orthotropous,  campylotropous,  and  anatropous,  and  the  modi- 
fication called  half-anatropous,  or  amphitropous.* 

521.  Orthotropous  (Fig.  580,  582,  585),  or  straight  o^ule,  is 
the  simplest  but  least  common  species,  being  that  in  which  the 


chalaza  is  at  the  evident  base,  and  the  orifice  at  the  opposite 
extremity,  the  whole  ovule  straight  (as  the  first  part  of  the  name 
denotes)  and  symmetrical.  Atropous,  meaning  not  turned  at 
all,  is  a  later  and  etymologically  much  better  name,  but  it  has 

1  An  epidermal  stratum  or  tegument  may  not  be  wanting  to  such  ovules, 
forming  a  sort  of  adherent  covering ;  but  this  in  nature  and  origin  is  not 
similar  to  the  ovular  coats. 

2  In  Latin  form,  orthotropa,  campylotropa,  anatropa,  amphitropa,  —  names  given 
by  Mirbel,  and  referring  to  the  way  in  which  the  ovule  is  turned  either  on 
itself  or  on  its  support.     Some  English  botanists  incongruously  write  ortho- 
tropal,  campylotropal,  &c. 

PIG.  685.  Orthotropous  or  Atropous  ovule  of  Buckwheat.  586.  Campylotropons 
ovule  of  Chickweed.  587.  Amphitropous  ovule  of  Mallow.  588  Anatropous  ovule  of  a 
Violet.  The  letter  h  indicates  the  hilum ;  c,  the  chalaza,  which  in  585  and  586  corre- 
sponds to  the  hilum;  /,  the  foramen  or  orifice;  r,  the  rhaphe. 


OVULES.  279 

not  come  into  general  use.     This  ovule  is  characteristic  of 
Polygonacese,  the  proper  Urticaceae,  Cistaceae,  &c. 

522.  Campylotropous  (Fig.  583,  586)  is  the  name  of  the  ovule 
which  in  the  course  of  its  growth  is  curyed  on  itself  so  as  to 
bring  the  orifice  or  true  apex  down  close  to  the  base,  here  both 
chalaza  and   hilum.      This  and  the  orthotropous  ovule  begin 
their  development  on  the  placenta  in  the  same  way,  but  the 
campylotropous  develops  unequally,   one  side  enlarging  much 
more   than   the  other,  especially  at   the  base,  until  the  ovule 
becomes   reniform,  and  chalaza   and   orifice   are  brought   into 
close  proximity.     Campylotropous  ovules  are  characteristic  of 
Cruciferae,    Capparidaceae,    Resedaceae,    Caryophyllaceae,    and 
Chenopodiacese. 

523.  Amphitropous  (Fig.  584,  587),  also  termed  ffeterotropous 
and  sometimes  Half -anatropous,  is  between  the  preceding  and 
the  following  ;  and  it  passes  in  various  instances  either  into  the 
one  or  into  the  other.     The  body  of  the  ovule  is  straight  or 
straightish,  but  it  stands  as  it  were  transversely  or  at  right 
angles  to  the  funiculus  and  hilum ;  and  it  is  fixed  by  the  middle, 
the  chalaza  at  one  end,  the  orifice  at  the  other.     An  apparent 
continuation  of  the  funiculus,  adherent  to  the  outer  coat,  extends 
from  the  hilum  to  the  chalaza.     Compared  with  the  preceding 
form,  the  explanation  is,  that  the  unequal  development  at  ite 
formation  is  confined  to  the  basal  half,  and  the  axis  remains 
straight,  while  the  whole  is  half  inverted  by  the  very  unequal 
growth.      Compared  with  the  next  form,  the  inversion  is  less 
and  the  later  growth  or  extension  of  the  apical  portion  greater. 
The  amphitropous  ovule  is  characteristic  of  Primulaceae,  and 
is  common  in  Leguminosaa. 

524.  Anatropous  (Fig.  588,  also  579,  581,  597)  is  the  name 
of  far  the  commonest  species  of  ovule,  that  in  which  the  organ, 
under  the  course  of  its  growth,  is  quite  inverted  on  its  base  ;  so 
that,  instead  of  standing  at  right  angles  with  the  funiculus,  it  is 
parallel  with  it,  or  rather  with  the   apparent  continuation  of 
it,  which  is  adherent  to  its  surface  as  a  sort  of  ridge  or  cord 
extending  along  the  whole  length  of  the  ovule,  from  hilum  to 
chalaza.     The  latter  occupies  the  seeming  apex  of  the  seed ; 
and  the  organic  apex  or  orifice  is  at  the  other  end,  close  beside 
the  hilum.     At  maturity,  the  ovule  is  straight,  but  not  wholly 
symmetrical,  the  attachment  being  oblique  or  somewhat  lateral, 
and  the  ridge  or  cord  on  that  side  not  rarely  prominent. 

525.  The  cord  or  ridge,  which  extends  along  the  whole  length 
of  the  anatropous  ovule,  and  for  half  its  length  in  the  amphi- 
tropous (Fig.  588,  587,  r),  is  named  the  RHAPHE.     This  is  not 


280  THE   FLOWER. 

at  all  a  seam,  as  the  Greek  word  denotes.  Its  origin,  and  the 
whole  structure  of  such  ovules  will  be  apprehended  by  comparing 
various  stages  of  its  growth. 

526.  Ap  ovule  of  any  kind  at  the  beginning  is  an 
excrescence  or  outgrowth  of  the  placenta,  or  of  some 
part  of  the  leaf-surface  if  there  is  no  developed 
placenta.  This  incipient  ovule  is  the  nucleus  (518), 
or  the  nucleus  surmounting  a  rudimentary  funiculus. 
The  nucleus  is  soft  cellular  tissue  only,  from  first  to 
last.  The  achlamydeous  ovule  (519)  undergoes  no 
further  development  except  in  size  or  shape.  Indeed 
sometimes  (as  in  Balanophorese)  this  bare  nucleus 
is  reduced  to  a  few  cells  of  parenchyma. 

527.  In  ordinary  ovules  a  new  growth  early  begins  around 
the  base  of  the  nucleus,  or  is  sometimes  coetaneous  with  it,  at 
first  as  a  ring  (or  part  of  a  ring) ,  soon  as  a  cup,  at  length  as 
an  enclosing  sac  or  covering,  open  at  the  top  ;  this  is  the  inner 
coat  of  the  ovule  when  there  are  two.    The  outer  coat  begins  and 
goes  on  in  the  same  way,  and  at  length  grows  over  and  encloses 
the  inner  coat  as  that  did  the  nucleus.     (Fig.  590-595.)    When- 
ever there  is  a  third  and  more  exterior  coat  it  is  formed  during 
the  growth  of  the  fertilized  ovule  into  the  seed,  to  which  there- 
fore it  belongs,  and  in  which  it  takes  the  name  of  arillus.    (597.) 
At  the  time  of  fertilization  the  apex  of  the  nucleus,  or  a  pro- 
longation of  it,  usually  projects  beyond  the  orifice  and  there 
receives  the  descending  pollen-tube.    Some  fibro- vascular  tissue, 
especially  spiral  ducts,  may  be  found  in  the  funiculus  and  cha- 
laza,  sometimes  extending  into  the  coats. 

528.  The  development  of  the  orthotropous  or  atropous  (un- 
turned) ovule  proceeds  symmetrically,   without  distortion,  the 


parts  keeping  their  primitive  direction.     In  the  campy lotropous, 
the  whole  of  one  side  of  the  ovule  greatly  outgrows  the  other. 

FIG.  689.  Magnified  view  of  a  vertical  section  of  a  carpel  of  Magnolia  Umbrella, 
about  a  month  before  anthesis,  showing  one  of  the  two  nascent  ovules,  at  this  time 
only  nucleus. 

FIG.  590-597.  Further  development  of  the  ovule  of  Magnolia  Umbrella,  showing  the 
formation  of  the  coats  and  the  anatropy.  590.  Ovule  a  week  older  than  in  589.  591. 
Same  a  week  or  two  later.  592.  Same  a  few  days  later.  693.  Same  from  a  nearly  full- 
grown  flower-bud.  594.  Sameattimeof  anthesis.  595.  Vertical  section  of  the  last  through 
the  middle  of  the  rhaphe.  596.  Cross-section  of  the  same.  (See  Jour.  Linn.  Soc.  ii.  108.) 


OVULES.  281 

In  the  anatropous,  the  inequality  of  growth  is  mainly  confined 
to  the  base  or  chalazal  region,  which  ends  by  becoming  upper- 
most ;  and  the  full-grown  ovule  has  the  ap- 
pearance of  being  inverted  on  and  adherent 
to  the  upper  portion  of  its  funiculus,  the 
rhaphe.  Fig.  589-597  illustrate  the  course  d 
of  development  from  a  comparatively  early 
period. 

529.  The  direction  of  anatropy  or  of  other 
turning  of  the  ovule  in  the  course  of  growth 
is  somewhat  diverse.  But  in  general,  when- 
ever ovules  are  in  pairs,  the  two  turn  from 
each  other,  in  the  manner  of  Fig.  315,  and  so  present  their 
rhaphes  back  to  back.  The  rhaphe-bearing  may  therefore  be 
called  the  dorsal  side  of  the  anatropous  ovule.  The  same  is  true 
in  the  case  of  numerous  ovules,  viz.,  those  of  one  half  of  the 
placenta  (or  one  leaf-margin)  turn  their  backs  to  those  of  the 
other.  When  such  ovules  are  solitary  or  in  single  rows,  and 
either  ascending  or  hanging,  the  rhaphe  is  usually  on  the  side 
next  to  the  placenta  or  ventral  suture,  as  in  Fig.  579  :  it  is  then 
said  to  be  ventral  (i.  e.,  next  the  ventral  suture),  or  adverse  to 
the  placenta.  In  certain  cases,  mostly  in  hanging  ovules,  as  in 
Fig.  581,  the  rhaphe  looks  in  the  opposite  direction,  toward  the 
dorsal  suture  or  midrib  of  a  simple  ovary :  it  is  then  said  to  be 
dorsal  or  averse  from  the  placenta.1 

1  By  comparison  of  Fig.  578  with  576  and  the  like,  it  may  be  perceived 
that  the  difference  is  explicable  by  a  kind  of  resupination  of  the  ovule  of 
the  former.  That  of  Ranunculus,  if  inserted  higher,  would  become  hori- 
zontal ;  and  if  the  insertion  were  transferred  to  the  very  summit  of  the  cell, 
it  would  be  suspended  and  the  rhaphe  averse,  as  in  Fig.  581.  Upon  this 
conception,  Euphorbia  and  its  allies  has  normally  suspended  ovules,  the 
rhaphe  being  next  the  placental  axis,  and  Buxus  and  its  allies,  resupinately 
suspended  ovules,  the  rhaphe  averse.  The  propriety  of  regarding  the  ad- 
verse rhaphe  as  the  normal  condition  is  confirmed  by  the  fact  that  the  only 
instance  we  know  of  solitary  erect  ovules  from  the  base  of  the  cell  having 
the  rhaphe  averse  is  that  of  Rhamnus  and  its  allies ;  and  here  it  was  shown 
by  Bennett  (in  PI.  Javan.  Rar.  131),  and  confirmed  by  the  analyses  of 
Sprague  (Gray,  Gen.  111.  ii.  168,  plates  163-169),  that  the  rhaphe  of  the  young 
ovules  is  ventral,  so  that  the  dorsal  position,  when  it  occurs,  is  the  result 
of  torsion.  J.  G.  Agardh  (in  his  Theor.  Syst.  PI.  178,  &c.)  maintains  the 
contrary,  but  is  not  sustained  by  later  observers. 

Accordingly,  even  if  we  adopted  Agardh's  estimate  of  the  botanical 
value  of  the  characters  here  considered,  we  should  prefer  to  express  these 
differences  in  the  phraseology  above  indicated,  and  not  to  adopt  his  terms, 

FIG.  597.  Same  as  595  more  magnified;  the  outer  coat,  (a),  the  inner  (b),  nucleus  (c), 
and  the  bundle  of  spiral  ducts  (d)  in  the  rhaphe  (running  from  placenta  to  chalaza) 
indicated. 


282  THE  FLOWER. 

530.  Origin  and  Nature  of  the  Ovule.     It  has  been  already 
stated  in   general   terms   that   ovules   are   peculiar  outgrowths 
or  productions,  generally  of  the  margins  of  carpellary  leaves 
(515)  ;  that  they  are  composed  of  parenchymatous  cellular  sub- 
stance, at  least  as  to  the  nucleus,  of  which  the  simplest  ovule 
wholly  consists  (526)  ;  that  the  coats  originate  subsequent!}-  to 
the  nucleus  ;  and  that  the  outer  coat  is  of  later  origin  than  the 
inner  one.   (518.)  The  mamiliform  protuberance  of  which  the 
forming  ovule  at  first  consists  originates  in  one  or  more  cells  of 
a  layer  directly  beneath  the  epidermis.1 

531.  The  morphological  nature  of  the  ovule  has  been  much 
discussed.     The  commonly  prevalent  view  was  that  the  ovule 
is  homologous  with  a  leaf-bud,  and  that  its  nature  is  in  some 
degree  illustrated  by  such  buds  as  those  which  develop  on  the 
margins  of  the  leaves  of  Bryophyllum,  as  shown  in  Fig.  322. 
But  such  buds,  and  the  bulblets  or  fleshy  buds  which  appear  on 
the  face  of  certain  leaves,  follow  the  universal  order  of  budding 
growth,  that  is,  are  centripetal  in  development,  the  outermost 
parts  being  the  earlier  and  the  inmost  the  later  formed.     The 
ovule,  on  the  contrary,  is  basipetal  or  centrifugal  in  develop- 
ment, the  nucleus  being  first  and  the  outer  coat  last  formed ; 
therefore  the  coats  are  not  homologous  with  sheathing  leaves, 
nor  the  nucleus  with  a  vegetative  axis.     The  older  theory  has 
accordingly  given  way  to  the  present  one,  in  which  the  ovule 
answers  to  the  lobe  of  a  leaf  peculiarly  transformed,  or  to  an 
outgrowth  of  a  leaf,  whether  from  its  edges  or  surface.     The 


apotropoas,  epitropous,  and  heterotropous  (the  first  two  new,  the  last  employed 
in  a  new  sense),  the  more  so  since  the  application  is  confused  with  hypo- 
thetical considerations  and  the  necessity  of  bringing  the  ovules  ideally  back 
to  ascending  or  horizontal  positions.  It  may  be  stated,  briefly,  that  Hetero- 
tropous, in  Agardh's  terminology,  applies  to  the  normal  position  of  collat- 
eral ovules,  with  rhaphes  back  to  back,  in  opposite  directions  on  the  two 
halves  of  the  placenta ;  Apotropous,  to  an  erect  or  ascending  ovule  with  its 
rhaphe  next  the  placenta!  axis,  and  a  hanging  one  has  its  rhaphe  averse 
from  it ;  Epitropous,  when  an  erect  or  ascending  ovule  has  its  rhaphe  averse, 
and  a  hanging  one  has  it  adverse. 

1  Hofmeister's  statement  that  the  simple  ovule  of  Orchis  originates  in 
the  division  of  a  single  epidermal  cell  (and  is  therefore  a  trichome)  is  con- 
troverted by  Strasburger  and  by  Warming.  The  latter  adds  the  remark, 
that  even  if  it  were  so  in  cases  of  extreme  simplicity,  this  would  not  invali- 
date the  proposition  that  the  ovule  is  to  be  regarded  as  the  homologue  of  the 
lobe  of  a  leaf.  Such  a  lobe  is  not  rarely  reduced  to  a  single  bristle.  For 
the  whole  subject  of  the  origin,  development,  morphology,  and  theory  of  the 
ovule,  see  Warming's  very  elaborate  and  perspicuous  memoir,  De  1'Ovule ; 
also  the  papers  of  Celakowsky,  Van  Tieghem,  &c.,  referred  to  in  notes  to 
paragraphs  500,  601. 


OVULES. 


283 


great  advantage  of  this  view  is  that  it  serves  to  homologize  the 
fructification  of  Flowering  Plants  with  that  of  the  higher  Flower- 
less  Plants,  or  the  Ferns,  the  sporangia  or  analogues  of  the 
ovule  being  outgrowths  of  the  leaf.1 

532.  Origination  of  the  Embryo.  The  whole  process  of  fer- 
tilization and  the  resulting  produc- 
tion of  the  embryo,  also  the  history 
of  the  subject,  belongs  to  the  suc- 
ceeding volume,  involving  as  they 
do  questions  of  minute  anatomy  and 
of  physiology.  But  a  general  idea 
may  here  be  given  of  the  way  in 
which  the  embryo  originates.  The 
tube  which  a  grain  of  pollen  sends 
forth  into  the  stigma  (574,  575) 
penetrates  the  style  through  loose 
conducting  tissue  charged  with 
nourishing  liquid,  reaches  the  cavity 
of  the  ovary,  enters  the  orifice  of  an 
ovule  to  reach  the  apex  of  the  nu- 
cleus, although  the  latter  sometimes 
projects  to  meet  the  pollen-tube. 
Meanwhile  a  cavity  (the  embryo- 
sac,  which  is  formed  by  the  great 
enlargement  of  a  single  cell  of  the 
tissue,  or  of  two  or  more  cells  the 
product  of  a  mother  cell)  forms  in 
the  nucleus,  the  upper  part  of  it 
commonly  reaching  nearly  or  quite 
to  the  apex  of  the  nucleus,  which 
the  pollen-tube  impinges  on  or 
sometimes  penetrates.  A  particular  portion  of  the  protoplasm 
contained  in  the  embryo-sac  forms  a  globule,  and  this  at  the  time 

1  The  advocates  of  this  view  naturally  maintain  that  ovules  and  placentae 
always  belong  to  leaves,  and  never  truly  to  a  cauline  axis ;  that  in  the  pre 
central  placentation  of  Primulaceae,  the  actual  ovuliferous  surface  is  an  out- 
growth of  the  bases  of  the  carpellary  leaves  coalescent  with  each  other  and 
adnate  to  a  prolongation  of  the  torus ;  also  that  in  those  Gymnosperms  which 
have  no  carpophyll,  such  as  Yew,  the  whole  nascent  carpellary  leaf,  or  rather 
the  papilla  which  would  otherwise  develop  as  such,  is  directly  developed  into 
ovule.  This,  being  solitary  and  the  last  production  of  the  axis,  necessarily 
appears  to  terminate  it.  (500,  601,  notes.) 

FIG.  598.  Diagram  representing  a  magnified  pistil  of  Buckwheat,  with  longitudinal 
section  through  the  axis  of  the  ovary  and  orthotropous  ovule ;  some  pollen  on  the  stigmas, 
one  grain  distinctly  showing  its  tube,  which  has  penetrated  the  style,  reappeared  in  the 
cavity  of  the  ovary,  entered  the  mouth  of  the  solitary  ovule  (o),  and  reached  the 
«nbryo-sac  (s)  near  the  embryonal  vesicle  (v). 


284 


THE   FLOWER. 


of  fertilization  is  found  at  the  apex  of  the  sac,  at  or  adjacent  to 

the  part  reached  by  the  pollen- tube.    Not  rarely  it  adheres  to  the 

599          era  eol  602         BOS  wall  of  the  sac  exactly 

opposite  the  termination 
of  the  pollen-tube.     This 
is    called    the    embryonal 
vesicle.     To  it   the    con- 
tents of  the  pollen-tube 
are  in  some  manner  trans- 
ferred.     Upon   which   it 
takes    a    more     definite 
shape,  acquires  a  wall  of 
cellulose,  and  so  becomes 
a   vegetable    cell.      This 
divides  into  two,  the  lower 
again  into  two,  and  so  on,  forming  a  chain  (the  suspensor  or  pro- 
embryo).     The  terminal  cell  of  this  divides  again  and  again  in 
three  directions,  producing  a  mass  of  cells  which  shapes  itself 
604          60s  606  607  into  the  embryo,  the  initial 

af\  t\        /HSv/l  plant  of  a  new  Senera- 

/I         d  Ik    If     JlHH    tion'    OrdinarilJ the  sus- 
pensor soon   disappears. 

It  is  attached  to  the  ra- 
dicular  end  of  the  em- 
bryo, which  consequently  always  points  to  the  foramen  or 
micropyle  of  the  seed.  The  process  in  Gymnosperms  is  more 
complex,  and  has  to  be  separately  described. 

533.  Polyembryony,  the  production  of  two  or  more  embryos  in 
one  seed,  is  not  uncommon  in  Gymnosperms  (there  being  a  kind 
of  provision  for  it),  and  is  of  occasional  but  abnormal  occurrence 
in  Angiosperms,  in  the  seed  of  Mistletoe,  Santalum,  &c.  In 
these  it  results  from  the  production  and  fertilization  of  more 
than  one  embryonal  vesicle.  Strasburger  has  recently  ascer- 
tained that  the  commoner  polyembryony  in  the  seeds  of  Onions, 
Oranges,  Funkia,  &c.,  results  from  the  production  of  adventive 
embryos,  which  originate  in  the  nucleus  outside  of  the  embryo- 
sac  and  wholly  independent  of  fertilization.1  Two  kinds  of 

1  Strasburger,  Ueber  Polyembryonie,  in  Zeitschr.  Naturwis.  Jena,  xii. 
1878  (see  Amer.  Jour.  Sci.  April,  1879).  It  was  found  that  when,  by  exclu- 
sion of  pollen,  the  formation  of  a  normal  embryo  was  prevented,  no  adventive 

FIG.  599.  Diagram  of  the  suspensor  and  Incipient  embryo  at  its  extremity.  600. 
The  same,  with  the  embryo  a  little  more  developed.  601.  The  same,  more  developed 
still,  the  cotyledons  faintly  indicated  at  the  lower  end.  602.  Same,  with  the  incipient 
cotyledons  more  manifest.  603.  The  embryo  nearly  completed. 

FIG.  604-606.  Forming  embryo  from  a  half-grown  seed  of  Buckwheat,  in  three  stages. 
607.  Same,  with  the  cotyledons  fully  developed. 


THE  FRUIT.  285 

anomalous  reproduction  are  therefore  now  known,  which  are 
intermediate  between  sexual  and  non-sexual,  between  budding 
and  fruiting  propagation,  viz., — 

Apogamy,  which  is  budding  growth  or  prolification  in  place  of 
that  which  should  subserve  sexual  reproduction.  This  was  dis- 
covered in  F«*rns  by  Prof.  Farlow,  while  a  pupil  of  De  Bary,  by 
whom  our  knowledge  of  the  process  has  recently  been  extended, 
and  this  nane  imposed.1  The  production  of  bulblets  in  place  of 
seed  or  embryo  answers  to  this  in  Flowering  plants. 

Parthen  >geny,  the  counterpart  analogue  of  apogamy,  is  the 
non-sexual  origination  of  an  embryo  extraneous  to  the  embryonal 
vesicle  rr  even  the  embryo-sac.  However  abnormal,  its  occur- 
red* u  probably  not  so  rare  as  has  been  supposed. 


CHAPTER  VII. 

THE  FRUIT. 

SECTION  I.   ITS  STRUCTURE,  TRANSFORMATIONS,  AND  DEHISCENCE. 

534.  The  Fruit  consists  of  the  matured  pistil  or  gynoecium 
(as  the  case  may  be) ,  including  also  whatsoever  may  be  joined 
to  it.  It  is  a  somewhat  loose  and  multifarious  term,  applicable 
alike  to  a  matured  ovary,  to  a  cluster  of  such  ovaries,  at  least 
when  somewhat  coherent,  to  a  ripened  ovary  with  calyx  and 
other  floral  parts  adnate  to  it,  and  even  to  a  ripened  inflores- 
cence when  the  parts  are  consolidated  or  compacted.  Fruits, 
accordingly,  are  of  various  degrees  of  simplicity  or  complexity, 
and  should  be  first  studied  in  the  simpler  forms,  namely,  those 
which  have  resulted  from  a  single  pistil.  Such  a  fruit  consists 
of  Pericarp  with  whatever  may  be  contained  in  it  and  incorpo- 
rated with  it. 


embryo  appeared  in  those  seeds  which  habitually  produce  them.  To  this 
Caelebogyne  offers  an  exception.  The  female  of  this  dioecious  plant  habit- 
ually matures  fertile  seeds,  with  a  well-formed  embryo,  in  Europe  when  there 
are  no  male  plants  in  the  country.  Strasburger  ascertained  that  the  embryo 
thus  formed  is  adventive,  the  embryonal  vesicle  perishing.  Parthenogenesis, 
of  which  Caelebogyne  was  the  most  unequivocal  case,  is  thus  confirmed,  and 
is  shown  to  occur  in  most  polyembryony ;  but  it  is  at  the  same  time  explained 
to  be  a  kind  of  prolification. 
*  See  Farlow,  in  Proc.  Am.  Acad.  ix.  68;  De  Bary,  Bot.  Zeit.  xxxvi.  466-487. 


286  THE  FRUIT. 

535.  The  Pericarp,  or  Seed-vessel,  is  the  ripened  ovary.     It 
should,  therefore,  accord  in  structure  with  the  ovary  from  which 
it  is  derived.     Yet  alterations  sometimes  take  place  during  fruc- 
tification, either  by  the  abortion  or  obliteration  of  parts,  or  by 
accessory  growth. 

536.  Internal  Alterations.     Thus,  the  ovary  of  the  Oak  con- 
sists of  three  cells,  with  a  pair  of  ovules  in  each  ;  but  the  fruit 
has  a  single  cell,  filled  with  a  solitary  seed,  only  one  ovule  being 
matured,  while  two  cells  and  five  ovules  are  suppressed,  the 
remains  of  which  may  be  detected  in  the  acorn.     The  ovary 
of  the  Chestnut  has  six  or  seven  cells,  and  a  pair  of  suspended 
ovules  in  each ;   but  only  one  of  the  dozen  or  fourteen  ovules 
ever  develops  into  a  seed,  except  as  a  rare  monstrosity.     The 
three-celled  ovary  of  the  Horsechestnut  and  Buckeye  is  similar 
in  structure    (Fig.    608-611),  and   seldom 

ripens  more  than  one  or  two  seeds ;  but  the 


abortive  seeds  and  cells  are  obvious  in  the  ripe  fruit.  The 
ovary  of  the  Birch  and  of  the  Elm  is  two-celled,  with  a  single 
ovule  in  each  cell :  the  fruit  is  one-celled,  with  a  solitary  seed ; 
one  of  the  ovules  being  uniformly  abortive,  while  the  other  in 
enlarging  thrusts  the  dissepiment  to  one  side,  and  obliterates  the 
empty  cell.  Similar  suppressions  in  the  fruit  of  parts  actually 
extant  in  the  ovary  are  not  uncommon. 

537.  On  the  other  hand,  there  may  be  more  cells  in  the  fruit 
than  there  are  primarily  in  the  ovary.  Thus  the  fruit  of  Datura 
is  dicarpellary  and  normally  two-celled,  with  a  large  placenta 
projecting  from  the  axis  far  into  the  cells.  But  each  cell  be- 
comes bilocellate,  that  is,  divided  into  two,  by  a  false  partition 
growing  out  from  the  back  of  each  carpel  and  cohering  with  the 
middle  of  the  adjacent  placenta.  So  the  5-carpellary  and  nor- 
mally five-celled  ovary  of  common  Flax  early  becomes  spuriously 
ten-celled  (morphologically  speaking,  not  10-locular,  but  10- 
locellate),  by  a  false  partition  extending  from  the  back  of  each 

FIG.  608.  Longitudinal  section  of  the  ovary  of  a  Buckeye  (jEscnlus  Pavia),  showing 
the  pairs  of  ovules  in  two  of  the  cells.  609.  Transverse  section  of  the  same  displaying  all 
three  cells  and  six  ovules.  610.  Same  of  half-grown  fruit,  with  single  fertile  seed,  abor- 
tive ovuleg  and  obliterating  cells.  611.  Dehiscent  one-seeded  fruit,  diminished  in  size. 


ITS   STRUCTURE  AND   TRANSFORMATIONS.  287 

carpel  across  its  cell  (Fig.  539-541)  ;  and  the  solitary  carpel 
is  similarly  divided  lengthwise  in  many  species  of  Astragalus, 
as  in  Fig.  534.  Transverse  divisions  or  constrictions  across 
a  maturing  ovary  (such  as  is  seen  in  Fig.  620)  are  not  uncom- 
mon, especially  in  legumes  and  other  pods,  and  are  of  little  mor- 
phological significance. 

538.  External  Accessions  may  here  be  referred  to.     The  wing 
of  the  pericarp  in  Maple,  Ash,  and  the  like  (Fig.  625-627) ,  are 
familiar  instances  of  this ;    and  of  the  same  nature  are  the  im- 
bricated scales  which  cover  some  Palm-fruits ;  the  prickles  on 
the  pod  of  Datura,  Ricinus,   &c.,  and  the  hooked  or  barbed 
prickles  of  many  small  pericarps  (as  in  various  Borraginaceae) , 
which  thus  become  burs  and  are  disseminated  by  adhering  to  the 
hairy  coat  of  cattle.     All  these  are  of  the  nature  of  superficial 
outgrowths,  and  these  especially  affect  the  pericarp  or  parts 
connected  with  it. 

539.  Persistence  of  Connected  Organs.    An  adnate  calyx  (331) , 
being  consolidated  with  the  ovary,  necessarily  makes  a  constit- 
uent part  of  the  fruit,  in  the  pome  (575)  doubtless  a  very  large 
part.     The  limb  or  lobes  of  such  adnate  organ  may  persist,  as 
the  tips  of  the  sepals  on  an  apple  or  quince,  and  may  be  turned 
to  useful  account,  as  is  the  pappus  of  Composite  for  dissemina- 
tion.   Or,  in  small  pericarps,  the  style  may  persist  as  part  of  the 
fruit,  and  subserve  the  same  ends,  either  by  becoming  feathery 
for  aerial  dissemination,  as  in  Clematis  and  in  one  section  of 
Geum,  or  by  becoming  hooked  at  the  tip  for  adhesion  to  fleece, 
&c.,  as  in  other  species  of  the  latter  genus.     Or  adjacent  parts 
which  are  not  actually  incorporated  with  the  pericarp  may  play 
similar  parts  in  the  economy,  as  the  hooks  on  the  calyx- tube  of 
the  dry  calyx  of  Agrimonia,  which  at  maturity  is  detached  with 
the  included  fruit,  the  fleshy  fructiferous  calyx  of  Gaultheria  (Fig. 
651)  and  of  Mulberry  (Fig.  654)  ;  and  the  pulpy  fructiferous  re- 
ceptacle of  the  strawberry  (Fig.  653)  :  the  ultimate  utilities  in 
both  classes  of  instances  being  similar,  viz.,  wide  dispersion  of 
the  seed  by  animals,  whether  by  external  carriage,  or  by  being 
devoured  and  the  voided  seeds  of  flesh y  fruits  thus  disseminated. 

540.  Transformations   in   Consistence.     In   the   change   from 
ovary  to  mature  pericarp,  various  kinds  of  transformations  may 
take  place.     In  some  the  wall  of  the  ovary  remains  thin  and 
becomes  in  fruit  foliaceous  or  leaf-like,  as  in  a  pea-pod,  the 
carpels  of  Columbine,  and  Marsh  Marigold  (Caltha) ,  or  the  pod 
of  Colutea  or  Bladder  Senna.    In  others  it  thickens  and  becomes 
at  maturity  either  dry  throughout,  as  in  nuts  and  capsules  ;  or 
fleshy  or  pulpy  throughout,  as  in  berries ;  or  hard-rinded  with- 


288  THE  FRUIT. 

out  but  soft  within,  as  in  a  pepo  ;  or  fleshy  or  berry-like  without, 
but  indurated  within,  as  in  all  stone-fruits,  such  as  the  cherry 
and  peach. 

541.  When  the  walls  of  a  pericarp  consist  of  two  layers  of  dis- 
similar texture  (as  in  a  peach)  the  outer  layer  is  called  EXOCARP, 
the  inner  ENDOCARP,  these  terms  meaning  exterior  and  interior 
parts  of  a  fruit.    When  the  external  layer  is  a  comparatively  thin 
stratum  or  film,  it  is  sometimes  termed  the  EPICARP.    When  it  is 
fleshy  or  pulpy  it  is  named  S  ARCOCARP.    When  the  endocarp  within 
a  sarcocarp  is  hard  and  bony  or  crustaceous,  forming  a  shell  or 
stone,  this  is  termed  a  PUT  AMEN.   When  three  concentric  layers  are 
distinguishable  in  a  pericarp,  the  middle  one  is  called  MESOCARP. 

542.  Fruits  may  be  divided  into  two  kinds,  in  reference  to 
their  discharging  or  retaining  the  contained  seeds.     They  are 
dehiscent  when  they  open  regularly  to  this  end  ;  indehiscent  when 
they  remain  closed.     There  is  a  somewhat  intermediate  condi- 
tion, when  they  rupture  or  burst  irregularly,  as  in  Datura  Metel, 
&c.     Dry  pericarps  with  single  seeds  are  commonly  indehiscent ; 
those  with  several  or  many  seeds  mostly  dehiscent.     Seeds  pro- 
vided with  a  wing  or  coma  or  any  analogous  help  to  dispersion 
are  always  in  indehiscent  pericarps.     Permanently  fleshy  peri- 
carps are  indehiscent,  stone-fruits  as  well  as  berries.     But  in 
some  stone-fruits   (i.  e.,   with  indurated  endocarp  and  fleshy 
exocarp),  such  as  those  of  Almond  (Fig.  640)   and  Hickory, 
the  barely  fleshy  exocarp  or  sarcocarp  dries  or  hardens,  instead 
of  softening,  as  maturity  is  approached,  and  at  length  separates 
from  the  putamen  by  dehiscence. 

543.  Dehiscence,  the  opening  of  a  pericarp  for  the  discharge 
of  the  contained  seeds,  is  vgular  or  irregular;  or,  better,  is 
normal  and  abnormal.     For  most  of  the  abnormal  or  non-typical 
modes  are  as  determinate  and  uniform  in  occurrence  as  the  typi- 
cal modes.     A  good  English  name  for  dehiscent  pericarps  in 
general  is  that  of  POD. 

544.  Regular  or  normal  dehiscence  is  that  in  which  a  pericarp 
splits  vertically,  for  its  whole  or  a  part  of  its  length,  on  lines 
which  answer  to  sutures  or  junctions,  that  is,  along  lines  which 
correspond  to  the  margins  or  midribs  of  carpellary  leaves,  or  to 
the  lines  and  surfaces  (or  commissures)  of  coalescence  of  con- 
tiguous carpels.     The  pieces  into  which  a  pericarp  is  thus  sun- 
dered are  termed  VALVES. 

545.  The  normal  dehiscence  of  a  carpel  is  by  its  inner,  ven- 
tral, or  ovuliferous  suture,  that  is,  by  the  disjunction  of  the 
leaf-margins,  as  in  Fig.   618.     Its  only  other  line  of  normal 
dehiscence  is  by  the  opposite  or  dorsal  suture,  that  is,  down 


DEHISCENCE. 


289 


the  midrib.     Legumes  usually  dehisce  by  both  sutures  (as  in 
Fig.  619),  therefore  into  two  valves. 

546.  A  dehiscent  pericarp  formed  of  two  or  more  carpels  is 
called  a  CAPSULE.    The  two  leading  terms  descriptive  of  capsular 
dehiscence  were  based  upon  the  modes  of  opening  of  pericarps 
having  as  many  cells  as  carpels :  they  are  the  septicidal,  that  is, 
as  the  term  denotes,  cutting  through  the  septa  or  dissepiments ; 
and  the  loculicidal,  that  is,  cutting  into  the  loculi  or  cells. 

547.  Septicidal,  the  dehiscence  through  the  dissepiments,  is 
the  disjunction  of  a  pericarp  into  its  constituent  carpels,  these 
then  usually  themselves   dehiscing  down  their  ventral  suture, 

as  in  Fig.  612,  illustrated  by 

the  diagram,  Fig.  613.    Good 

examples  are  furnished  by  the 

Hypericum  Family  (the  pistil 

illustrated  in  Fig.  536,  537), 

where    the    placentae    which 

compose  the  axis  are  carried 

away  on  the  edges  of  the  par- 
titions or  introflexed  valves ; 

also  by  Rhododendron,  Kal- 

mia,  and  the  like,  in  which 

the  placentae  remain  combined 

into  a  column  in  the  axis  (the 

COLUMELLA  or  column) ,  from 

which  the  edges  of  the  valves 

break  away. 

548.    The    septicidal    dis- 
junction of  the  carpels  does , 
Onot  of  itself  open  the  cells. 
Such  separated  carpels  when 
613  one-seeded  not  rarely  remain 

closed,  as  in  Mallow,  Ver- 
bena, dec.  Or  when  dehiscent  they  may  open  both  by  the  ventral 
and  dorsal  sutures;  i.  e.,  the  pericarp  may  first  divide  into  its 
constituent  carpels,  and  then  each  carpel  break  up  into  half 
carpels,  as  in  Euphorbia. 

549.  Loculicidal,  the  dehiscence  into  the  loculaments,  loculi, 
or  cells  of  the  pericarp  (shown  in  Fig.  614,  and  the  diagram, 
615),  is  that  in  which  each  component  carpel  splits  down  its 


PIG.  612.  Septicidally  dehiscent  tricarpellary  capsule  of  Elodes  Virginica.  613.  Dia- 
gram of  septicidal  dehiscence. 

FIG.  614.  Loculicidally  dehiscent  tricarpellary  capsule  of  an  Iris,  divided  trans- 
versely at  the  middle.  615.  Diagram  of  loculicidal  dehiscence. 


290  THE  FRUIT. 

dorsal  suture,  as  in  Iris,  Hibiscus,  GEnothera,  &c.  In  this,  the 
dissepiments  remain  intact.  If  they  break  away  from  the  centre 
then  they  are  borne  on  the  middle  of  the  valves,  as  in  the  figures 
above  cited.  If  they  remain  coherent  in  the  axis  but  break  away 
from  the  valves,  the  result  is  one  form  of  what  is  called  — 

550.  Septifragal  dehiscence,   i.  e.,   a   breaking   away  of  the 
valves  from  the  septa  or  partitions,  as  shown  in  Fig.  616.     This 
represents  the  loculicidal  form  of  the  septifragal  mode,  which  is 
less  common  than  that  of  the  accompanying  diagram,  Fig.  617. 

Here  the  partitions  alternate 
with  the  valves  ;  that  is,  the 
dehiscence  of  the  pericarp  is 
of  the  septicidal  order,  as 
near  as  may  be,  but  the  par- 
titions do  not  split,  wherefore 
the  valves  break  away  at  the 
common  junction.  To  this 
the  term  marginicidal  has  been  applied.  It  occurs  in  the  2-3- 
carpellary  capsule  of  Ipomsea  (especially  in  the  common  Morning 
Glory) ,  in  the  5-carpellary  capsule  of  the  North  American  species 
of  Bergia ;  likewise  in  the  2-carpellary  pod  of  Cruciferse  (Fig. 
623),  with  a  difference  that  the  placentae  from  which  the  valves 
break  away  are  here  parietal  and  the  partition  is  abnormal. 

551.  The  terms  septicidal  and  loculicidal  apply  equally  in  plan, 
though  not  with  etymological  correctness,  to  one-celled  capsules 
with  either  parietal  (495)  or  free  central  (599)  placentae.    When 
the  dehiscence  is  of  the  septicidal  type  and  the  placentation  pari- 
etal, the  (half)  placentas  are  borne  on  the  margin  of  the  valves, 
as  in  the  Gentian  family  and  the  species  of  Hypericum  with  one- 
celled  capsule.     When  the  placentae  are  borne  on  the  middle 
of  the  valves,  as  in  Violets,  the  dehiscence  is  of  the  loculicidal 
type.     In  the  case  of  free  central  placentas  with  no  trace  of 
partitions,  the  character  of  the  dehiscence  may  usually  be  deter- 
mined by  the  position  of  the  styles  or  stigmas  relative  to  the 
valves. 

552.  Dehiscence  may  be  quite  normal  although  very  partial, 
as  when  confined  to  the  apex  of  the  capsule  of  Cerastium  and 
of  Primula,  and  even  to  the  pores  under  the  radiate  stigmas 
of  Poppy. 

553.  Irregular  or  abnormal  dehiscence  is  such  as  has  no  respect 
to  the  normal  sutures ;   as  where  the  dehiscence  is  transverse  ; 


FIG.  616.    Diagram  of  loculiddally  septifragal  dehiscence.  617.  Same  of  septicidally 
or  rather  marginicidally  septifragal  debiscence. 


ITS  KINDS. 

either  extending  part  way  round,  as  in  the  pod  of  Jeffersonia, 
or  completely  round,  so  that  the  upper  part  falls  off  like  an 
unhinged  lid.  This  circumscissile  dehiscence  occurs  in  many 
plants  of  widely  different  orders  ;  such,  for  example,  as  Purslane 
(Fig.  621),  genuine  Amaranths,  Plantain,  Pimpernel,  and  Hen- 
bane. In  other  cases,  as  in  Antirrhinum  (Snap-dragon)  and 
its  allies,  the  cells  burst  by  irregular  laceration  at  a  definite 
point,  and  discharge  the  seeds  through  the  ragged  perforation  ; 
or  one  or  more  neat  valvular  orifices  are  formed  on  some  parts 
of  the  wall,  as  in  Campanula. 


SECTION  II.  THE  KINDS  OF  FRUIT. 

554.  FRUITS  have  been  minutely  classified  and  named;1  but 
the  terms  in  ordinary  use  are  not  very  numerous.     A  rigorously 
exact  and  particular  classification,  discriminating  between  the 
fruits  derived  from  simple  and  from  compound  pistils,  or  between 
those  with  and  without  an  adnate  calyx,  is  too  recondite  and 
technical,   and  sometimes  too  hypothetical,   for  practical  pur- 
poses.    It  is  neither  convenient  nor  philosophical  to  give  a 
substantive  name  to  every  modification  of  the  same  organ.     For 
all  ordinary  purposes,   both  of  morphological  and   systematic 
botany,  it  will  suffice  to  characterize  the  principal  kinds  under 
the  four  classes  of — 

Simple  fruits,  those  which  result  from  the  ripening  of  a  single 
pistil ; 

•Aggregate,  those  of  a  cluster  of  carpels  of  one  flower  crowded 
into  a  mass ; 

Accessory  or  Anthocarpus,  where  the  principal  mass  consists 
of  the  surroundings'  or  support  of  either  a  simple  or  an  aggregate 
fruit; 

Multiple  or  Collective,  formed  by  the  union  or  compact  aggre- 
gation of  the  pistils  of  several  flowers,  or  of  more  than  one. 

555.  Simple  Fruits  may  be  distinguished,  upon  differences  of 
texture,  into  Dry  Fruits,  Stone  Fruits,  and  Baccate  Fruits ;  or, 
better,  into  Dry  and  Fleshy  ;  and  the  first  may  be  divided  into 


1  The  greater  part  of  the  forty-three  substantive  names  of  Desvaux's, 
and  even  of  the  thirty-six  of  Dumortier's  and  of  Lindley's  elaborate  classi- 
fications of  fruits  have  never  found  employment  in  systematic  botany,  and 
doubtless  never  will  be  used.  Yet  a  detailed  carpological  classification  has 
its  uses  for  the  student.  Among  the  more  recent  attempts  are  the  successive 
ones  of  Dickson,  McNab,  and  Masters.  See  Nature,  iv.  347  (also  hi  Trimen's 
Jour.  Bot.  1871,  310),  iv.  475,  and  v.  6. 


292 


THE  FKTJIT. 


dehiscent  and  indehiscent  kinds.1  Theoretically,  each  kind  may 
be  divided  into  those  of  a  simple  and  those  of  a  compound  pistil, 
and  some  would  make  the  primary  division  on  this  character. 
Some  also  would  separate  fruits  with  adnate  or  superior  calyx 
from  those  free  of  all  such  combination.  But  in  practice  these 
differences  can  seldom  be  indicated  by  substantative  names. 
The  name  of  berry  is  equally  applicable  to  the  fruit  resulting 
from  the  single  carpel  of  Actsea,  the  syncarpous  ovary  of  the 
grape,  and  the  similar  ovary  with  adnate  calyx  of  a  gooseberry 
and  cranberry.  It  should  be  understood  that  the  kinds  shade 
off  one  into  another  most  freely. 

556.  Dehiscent  Fruits  (543),  or  Pods,  are  distinguishable  into 
apocarpous,  or  of  single  carpels,  and  syncarpous,  of  more  than 
one  carpel,  i.  e.  the  first  of  a  simple,  the  second  of  a  compound 
pistil.  The  first  kind  is  mainly  represented  by  the  Follicle  and 
the  Legume  ;  the  second,  by  the  Capsule  and  its  modifications. 
557.  A  Follicle  is  a  pod  formed 

of  a  simple  pistil,  and  dehiscent  by 

one   suture    (this   almost    always 

the  ventral  or  inner  suture)  alone  ; 

as   in   the   Larkspur,  Columbine, 

Peony,  and  Marsh-Marigold  (Fig. 

618)  ;  also  in  Milkweed  and  Dog- 
bane. There  may  be  several  follicles  or  only 
one  to  a  flower,  even  in  the  same  genus,  as  in 
Larkspurs,  Cimicifuga,  &c.  In  Magnolia 
(Fig.  648-650),  fleshy  carpels  become  follicles 
dehiscent  by  the  dorsal  suture. 

558.  A  Legume  is  the  pod  formed  of  a 
simple  pistil  which  is  dehiscent  by  both  sut- 
ures (as  in  the  Pea,  Fig.  619),  so  dividing 
into  two  pieces  or  valves.  (544.)  This  is  the 
fruit  of  the  Pulse  Family,  accordingly  named 
Leguminosse  (Leguminous  plants) :  indeed, 
the  name  of  legume  is  restricted  to  the  fruits  of  this  family, 
and  in  descriptive  botany  is  extended  to  all  the  modifications 


1  Dr.  Masters's  modification  of  Dickson's  and  McNab's  classification  of 
simple  fruits,  as  to  primary  kinds,  is  into 

1.  Nuts,  or  Achcenocarps,  dry  and  indehiscent ; 

2.  Pods,  or  Regmacarps,  dry,  dehiscent ; 

3.  Stone-fruits,  orPyrenocarps,  fleshy  without,  indurated  within,  indehiscent; 

4.  Berries,  or  Sarcocarps,  fleshy  throughout,  indehiscent. 

PIG.  618.    A  dehiscent  follicle  of  Marsh-Marigold,  Caltha  palustris. 

FIG.  619.    Legume  of  a  Sweet  Pea,  already  dehiscent.  620.  Loment  of  a  Desmodlum. 


ITS   KINDS. 


293 


which  that  order  presents.  Some  of  these,  in  fact,  are  in- 
dehiscent  and  reduced  to  akenes ;  some  break  up  at  maturity 
into  one-seeded  indehiscent  articulations  or  joints,  which  are 
dispersed  as  if  they  were  so  many  seeds.  A  legume  of  the  latter 
kind  takes  the  special  name  of  LOMENT,  Lat.  Lomentum.  (Fig. 
620.)  In  Mimosa  (Sensitive-plant,  &c.),  such  articulations  de- 
hisce into  two  valves.  They  also  fall  away  from  the  sutures, 
or  from  a  persistent  marginal  border  of  them,  or  in  some  cases 
the  valves  thus  fall  away  entire.  The  persistent  frame  which 
remains  has  been  called  a  REPLUM,  an  architectural  word,  here 
taken  in  the  sense  of  door-case. 

559.  A  Capsule  is  the  pod,  or  dehiscent  fruit,  of  any  compound 
pistil.      When    regularly   and    com- 
pletely dehiscent,   as  already  stated 

(544),  the  pod  splits  lengthwise  into 
pieces  or  valves.  The  modes  of  regular 
dehiscence  are  illus- 
trated in  Fig.  612- 
617.  Two  modifica- 
tions of  the  capsule 
have  received  distinc- 
tive names  which  are 
in  common  use,  viz. 
the  Pyxis  and  the 
Silique. 

560.  A  Pyxis  or  Pyxidium  is  a  dry  fruit  which  opens  by  a 
circular  line,  cutting  off  the  upper  part  as  a  lid  ;  i.  e.,  the  dehiscence 
is  circumscissile.  (553,  Fig.  621.)  In  the  Purslane, 

Pimpernel,  Henbane,  and  Plantain,  the  pyxis  is 
a  capsule ;  in  Amaranths  (Fig.  637)  it  is  a 
utricle ;  in  Jeffersonia  (Fig.  622)  it  is  a  modi- 
fication of  the  follicle,  being  of  one  carpel  which 
dehisces  transversely,  and  not  all  round,  so  that 
the  lid  remains  attached. 

561.  A  Silique  is  a  narrow  two-valved  capsule,  with  two  pari- 
etal placentae,  from  which  the  valves  separate  in  dehiscence  ;  as 
in   plants   of  the   Cruciferous   or  Mustard  family   (Fig.  623), 
to  the  fruit  of  which  this  term  is  restricted.     Usually,  a  false 
partition  is  stretched  across  between  the  two  placentae,  render- 

FIG.  621.  Pyxis  of  Purslane,  Portnlaca  oleracea,  the  top  separating  entirely  and 
falling  away. 

FIG.  622.  Pyxis-like  follicular  fruit  of  Jeffersonia  diphylla;  the  lid  remaining 
attached  doreally. 

FIG.  623.  Silique  of  Cardamine,  in  dehiscence.  624.  Silicle  of  Capsella  or  Shepherd'*- 
Purse,  lateral  view,  and  an  oblique  view  of  the  same  with  one  valve  removed. 


294 


THE  FET7IT. 


ing  the  pod  two-celled  in  an  anomalous  manner.  A  SILICLE 
(Silicula,  diminutive  of  siligua)  is  merely  a  short  silique,  the 
length  of  which  does  not  more  than  twice  or  thrice  surpass  the 
breadth;  such  as  that  of  Shepherd's-Purse  (Fig.  624),  and  of 
Lunaria,  Candytuft,  &c. 

562.  Indehiscent  Dry  Fruits  are  almost  always  one-seeded  or 
very  few-seeded.  If  numerous,  the  seeds  thus  placed  would  not 
be  dispersed.  The  ordinary  kinds  are  strictly  one- 
seeded,  and  in  common  language  are  often  con- 
founded with  seeds.  The  ways  in  which  such  fruits 
are  dispersed  are  various.  In  the  following  case, 
the  adaptation  of  the  pericarp  to  dispersion  by  wind 
distinguishes  the  species  of  fruit. 

563.  The  Samara,  sometimes  called  in  English  a 
Key,  is  an  indehiscent  one-seeded  fruit  provided 
with  a  wing.  In  the  White  Ash 
the  wing  is  terminal  (Fig.  625) ; 
in  other  species  the  whole  fruit 
is  wing-margined ;  in  Birch  and 
Elm  (Fig.  626)  the  wing  sur- 
rounds the  body  of  the  pericarp  ; 
and  the  Maple  fruit  is  a  double 
samara  or  pair  of  such  fruits,  con- 
626  spicuously  winged  from  the  apex. 
564.  Akene  (Lat.  Achenium)  is  a  general  name  for  all  the 
one-seeded,  dry  and  hard,  indehiscent  and  seed-like  small  fruits, 
such  as  are  popularly  taken  for  naked 
seeds.  But  that  they  are  true  pistils, 
or  ovaries  ripened,  is  evident  from  the 
style  or  stigma  they  bear,  or  from  the 
scar  left  by  its  fall ;  and  a  section 
brings  to  view  the  seed  within,  provi- 
ded with  its  own  proper  integuments. 
The  name  has  been  restricted  to  the  seed-like  fruits  of  simple 
pistils,  such  as  those  of  the  Buttercup  (Fig.  628, 629) ,  Anemone, 
Clematis,  and  Geum.  The  style  in  some  species  of  the  latter 
remains  on  the  fruit  as  a  long  and  feathery  tail,  in  others  as  a 
short  and  hooked  one,  both  being  agents  of  dissemination.  The 
grains  of  the  strawberry  (Fig.  653)  are  also  akenes.  The  name 
is  extended  to  all  one-celled  seed-like  fruits  resulting  from  a 

FIG.  625.  Samara  or  key  of  White  Ash,  Fraxlnus  Americana.  626.  That  of  White 
Elm.  Ulmus  Americana.  627.  Double  samara  of  Red  Maple,  Acer  rubrum. 

FIG.  628.  Achenium  of  a  common  Buttercup.  629.  Vertical  section,  showing  the 
seed  within. 


ITS   KINDS. 


295 


compound  ovary,  and  even  when  invested  with  an  adnate  calyx- 
tube.     Of  the  latter  is  the  fruit  of  Composite.    (Fig.  630-635.) 
Here  the  tube  of  the  calyx  is  incorpo- 
rated with  the  surface  of  the  ovary ; 
and  its  limb  or  border,  obsolete  in  some 
cases  (Fig.    630),  in   others   appears 
as  a  crown  or  cup  (Fig.  631),  or  set  of 
teeth  or  of  scales  (Fig.  632,  633),  or  as 
a  tuft  of  bristles  or  hairs  (Fig.  634, 


635),  &c.,  called  the  PAPPUS.  In  the  Lettuce  and  Dandelion 
(Fig.  635) ,  the  achenium  is  rostrate,  or  beaked,  i.  e.  its  summit 
is  extended  into  a  slender  beak.  An  akene  with  adnate  calyx 
has  been  termed  a  CYPSELA. 

565.  The  Utricle  is  the  same  as  the  akene,  only  with  a  thin 
and  bladdery  loose  pericarp,  like  that  of  Goosefoot. 

(Fig.  636.)  This  thin  coat  sometimes  bursts  irregu- 
larly, discharging  the  seed.  In  the  true  Amaranths, 
the  utricle  opens  by  a  circular  line,  and  the  upper 
part  falls  as  a  lid,  converting  the  fruit  into  a  small 
pyxis  (560),  —  a  transition  form.  (Fig.  637.) 

566.  A  Caryopsis  or  Grain  differs  from  the  utricle 
or  akene  in  having  the  seed  completely  filling  the 
cell,  and  its  thin  coat  firmly  consolidated  throughout 
with  the  very  thin  pericarp ;  as  in  wheat,  Indian 
corn,  and  all  other  cereal  grains.     Of  all  fruits  this 
is  the  kind  most  likely  to  be  mistaken  for  a  seed. 

567.  A  Jfut  is  a  hard,  one-celled  and  one-seeded,  indehiscent 
fruit,  like  an  achenium,  but  larger,  and  usually  produced  from 
an  ovary  of  two  or  more  cells  with  one  or  more  ovules  in  each, 
all  but  a  single  ovule  and  cell  having  disappeared  during  its 
growth  (536)  ;  as  in  the  Hazel,  Beech,  Oak  (Fig.  638),  Chest- 


FIG.  630.  Achenium  of  Mayweed  (no  pappus).  631.  That  of  Cichory  (its  pappus  a 
shallow  cup).  632.  Of  Sunflower  (pappus  of  two  deciduous  scales).  633.  OfSneezeweed 
(Helenium),  with  its  pappus  of  five  scales.  634.  Of  Sow-Thistle,  with  its  pappus  of 
delicate  downy  hairs.  635.  Of  the  Dandelion,  tapering  below  the  pappus  into  a 
long  beak. 

PIG.  636.  Utricle  of  Chenopodium  album,  or  common  Goosefoot.  637.  Utricle  of  an 
Amaranth,  by  transverse  dehiscence  becoming  a  pyxis. 


296  THE  FRUIT. 

nut,  and  the  like.     The  nut  is  often  enclosed  or  surrounded  by 
a  kind  of  involucre,  termed  a    Cupule;   such   as   the   cup   at 
the  base  of  the  acorn,  the  bur  of  the  chestnut,  and  the  leaf- 
like  covering  of  the  hazel-nut.     The  name  Glans 
(sometimes  Gland  in  English)  is  technically  applied 
to  such  nuts,  this  being  their  classical  Latin  name. 
568.   The  fruit  of  the  Walnuts  and  Hickory  is 
apparently  a  kind  of  drupaceous  nut,  or  something 
intermediate  between  a  stone  fruit  and  a  nut.     But 
certain  monstrosities  give  reason  for  supposing  that 
the    seeming    exocarp    (541),   which   in   Hickory 
638  hardens  and  at  maturity  dehisces  in  four  valves, 

is  of  the  nature  of  an  adnate  involucre.     The  cocoanut  is  a  sort 
of  fibro-drupaceous  nut. 

569.  Nutlet,  or  in  Latin  form  NUCULE  (Nucula),  is  sometimes 
superfluously  employed  in  a  literal  sense,  as  a  diminutive  nut.1 
Of  late  it  has  acquired  a  good  and  fairly  legitimate  use  as  the 
name  of  the  seed-like,  or  rather  akene-like,  closed  parts  or  lobes, 
of  crustaceous  or  other  hard  texture,  into  which  certain  bilocular 
or  plurilocular  pericarps  separate  at  maturity,  i.  e.  for  the  seg- 
ments of  a  schizocarp,  571,  which  resemble  akenes.2    These  are 
sometimes  carpels,  sometimes  half-carpels,  as  in  Verbena,  also 
in  Borraginaceae  and  Labiatae  (in  which  the  segments  are  greatly 
separated  in  the  ovary),  and  sometimes,  as  in  Nolana,  they  are 
portions  of  compounded  carpels  which  have  been  exceedingly 
multiplied  by  chorisis. 

570.  There  are  complete  transitions  between  dry  nutlets,  with 
a  thin  and  herbaceous  epicarp,  and  the  pyrence  (574)  or  stony 
inner  portion  of  such  carpels  when  drupaceous  or  composing  a 
drupe  of  two  or  more  stones.     It  is  therefore  a  hardly  incongru- 
ous and  very  convenient  use  which  extends  the  term  nutlet  to 
include  these  small  seed-like  stones  also,  as,  for  example,  to 
those  of  Holly,  Bearberry,  Hawthorn,  and  the  like. 

571.  The  pair  of  achenium-like  or  often  samara-like  carpels, 

1  Nut  and  akene,  between  which  there  is  no  fixed  distinction,  will  cover 
this  ground.     The  fruit  of  Cyperacese,  for  instance,  is  truly  an  achenium, 
if  this  name  is  ever  to  be  used  (and  it  now  commonly  is)  for  any  other  than 
a  monocarpellary  fruit.     It  is  often  termed  a  nut,  sometimes  a  nutlet,  and 
by  a  late  writer,  Boeckler,  a  caryopsis. 

2  Cocci  (sing.  Coccus,  from  a  Greek  word  for  kernel)  is  another  name  for 
fruit-carpels,  or  separating  lobes  of  a  dry  pericarp,  as  well  for  dehiscent  ones 
(of  Euphorbia)  as  for  indehiscent.     Hence  such   lobed  or  partible  fruits 
are  said  to  be  dicoccous,  tricoccous,  &c.,  according  to  the  number  of  lobes  or 
carpels. 

FIG.  638.    Acorn  (nnt)  of  White  Oak,  with  its  cup,  or  cnpule. 


ITS  KINDS. 


297 


united  by  their  inner  face  but  separating  entire  at  maturity, 
which  constitute  the  fruit  of  Umbelliferae,  takes  the  name  of 
CREMOCAKP  (Lat.  Cremocarpium)  ;  and  the  halves  are  called 
MERICARPS.  These  names  it  may  sometimes  be  convenient  to 
use ;  yet  it  is  not  advisable  to  have  special  names  for  the  fruits 
of  particular  families ;  and  mericarp  is  here  synonymous  with 
carpel.  For  dry  fruits  in  general  (or  such  as  become  dry) 
which  are  composed  of  two  or  more  carpels,  and  which  at  matu- 
rity split  up  or  otherwise  separate  into  two  or  more  closed  one- 
seeded  portions,  an  appropriate  recent  name  is  that  of  SCHIZOCABP. 
The  component  carpels  of  such  a  fruit  were  long  ago  named  Car- 
cerules  (carceruli,  little  prisons)  by  Mirbel. 

572.  Fleshy  Fruits,   which   from   their  texture  are  naturally 
indehiscent,  may  be  either  fleshy  throughout,  or  with  a  firm  rind 
or  shell,  or  fleshy  externally  and  hard  or  stony  internally.     Of 
the  latter,  the  t}-pe  is 

573.  The  Drupe  or  Stone  Fruit  proper  (Fig.  639),  that  of  the 
cherry,  plum,  and  peach.     True  drupes  are  of  a  single  carpel, 
one-celled    and    one-seeded 

(or  at  most  two-seeded) ,  in 

the  ripening  of  which   the      [ -ilj*  V^<  "¥  4  \     /         \V 
outer  portion  of  the  pericarp  s-  ^»  /  ^  V   i 

becomes  fleshy  or  pulpy,  and 
the  inner  stony  or  crustace- 
ous,  i.  e.  divides  into  sar co- 
carp  and  putamen.  (541.) 
But  the  name  is  extended  to 
pericarps  of  similar  texture  resulting  from  a  compound  pistil, 
either  of  a  single  cell,  as  in  Celtis,  and  (by  abortion)  in  the  olive, 
or  of  two  or  several  cells,  as  in  Cornus,  Khamnus,  &c.  The  several 
pericarps  of  the  aggregate  blackberry  and  raspberry  are  diminu- 
tive drupes  or  DRUPELETS. 

574.  Small  drupes  are  often  confounded  with  berries,  and  the 
stone  or  stones  taken  for  seeds.     Especially  is  it  so  in  drupes 
or  drupaceous  fruits  of  more  than  one  cell,  ripening  into  separate 
or  separable  hard  endocarps  or  stones,  each  filled  by  a  seed.1 
Bearberries  (Arctostaphylos)   and  Huckleberries  (Gaylussacia) 
are  good  illustrations  of  this.     The  seed-like  endocarps  of  this 

1  The  term  Acinus,  the  original  name  of  such  a  berry  as  a  grape,  has  been 
used  in  descriptive  botany  for  a  small  drupe  or  drupelet,  and  the  ripened 
carpels  of  Rubus  have  been  termed  acini  or  acines,  but  without  discriminating 
them  from  berries. 

PIG.  639.  Vertical  section  of  a  peach.  640.  An  almond ;  In  which  the  exocarp,  the 
portion  of  the  pericarp  that  represents  the  pulp  of  the  peach,  remains  juiceless,  and  at 
length  separates  by  dehiscence  from  the  endocarp,  or  shell 


298 


THE   FRUIT. 


sort  are  PYRENAE  ;  and  the  fruits  are  dipyrenous,  tripyrenous, 
tetrapyrenous,  &c.,  according  as  they  contain  two,  three,  or  four 
pyrenae.  When  the  sarcocarp  is  thin  and  dries  up  at  maturity, 
these  pyrenae  pass  by  gradations  into  nuculae  (569)  or  nutlets : 
hence  pyrenae  are  not  uncommonly  in  English  descriptions  called 
nutlets  or  nucules. 

575.  The  Pome  (Fig.  641,  642)  is  the  name  of  the  apple,  pear, 
and   quince.      These   are   fleshy   fruits,    composed   of   two   to 
several  carpels  (rarely  by  abortion  only  one) ,  of 
parchment-like  or  (in  Hawthorns)  bony  texture, 

enclosed  in  flesh  which  morphologically  belongs 
to  adnate  calyx  and  receptacle ;  as  may  be  ap- 
prehended by  comparing  a  rose-hip  (Fig.  407,  in 
flower)  with  an  apple  or  a  pear.  Of  the  quince,  the 
whole  flesh  is  calyx  or  hypanthium  (395)  ;  in  the 
apple  and  pear,  the  inner  or  core-portion  of  the 
flesh  is  of  the  nature  of  disk,  investing  the  carpels. 
In  the  fruit  of  Hawthorns,  the  carpels  become  bony 
pyrenae  (574),  and  so  the  fruit  is  drupaceous,  is 
indeed  nothing  more  than  a  syncarpous  drupe. 
In  Eriobotrya,  or  Cumquat,  the  carpels  becoming 
very  thin  and  membranaceous,  the  pomaceous 
fruit  is  in  fact  a  kind  of  berry. 

576.  The  Pepo,  or  Gourd-fruit  (Fig.  643),  of  which  the  gourd 
and  squash  are  the  type,  and  the  melon  and  cucumber  equally 

familiar  illustrations,  is  the  char- 
acteristic fruit  of  Cucurbitaceae, 
fleshy  internally  and  with  a  hard 
or  firm  rind,  nil  or  part  of  which 
is  referable  to  the  adnate  calyx 
completely  incorporate  with  the 
ovary.  This  is  either  one-celled 
with  three  broad  and  revolute 
parietal  placentae,  or  these  pla- 
centae, borne  on  thin  dissepiments, 
meet  in  the  axis,  enlarge,  and 
spread,  unite  with  their  fellows 
on  each  side,  and  are  reflected  to 
the  walls  of  the  pericarp,  next  which  they  bear  their  ovules.  As 
the  fruit  enlarges,  the  seed-bearing  placentae  usually  cohere 
with  the  walls,  and  the  partitions  are  obliterated,  giving  the 

FIG.  641.  Pome  or  apple  in  transverse  section.  642.  Quince  in  vertical  section:  the 
inner  flesh  answering  to  disk  in  the  apple  and  pear  is  here  wanting. 

FIG.  643.  Section  of  the  ovary  of  the  Gourd.  644.  Diagram  of  one  of  its  constituent 
carpel*. 


ITS   KINDS. 


299 


appearance  of  a  peculiar  abnormal  placentation,  which  the  study 
of  the  ovary  readily  explains.  In  the  watermelon  the  edible 
pulp  all  belongs  to  the  greatly  developed  placenta?.  Fruits  of 
this  family  in  which  the  rind  also  is  soft  at  maturity  are  true 
berries. 

577.  The  Hesperidium  (orange,  lemon,  and  lime)  is  the  fleshy 
fruit  of  a  free  many-celled  ovary  with  a  leathery  rind,  and  is  a 
mere  variety  of  the  berry.     The  name  is 

applied  only  to  fruits  of  the  Orange  tribe. 

578.  The  Berry  (Lat.  Bacca)  comprises 
all  simple  fruits  in  which  the  pericarp  is 
fleshj- throughout.    The  grape,  gooseberry, 
currant,  cranberry  (Fig.  645),  banana,  and 
tomato  are  familiar  examples.     The  first 
and  last  consist  of  an  ovary  free  from  the 
calyx ;  in  the  others,  calyx  and  ovary  are 
combined  by  adnation. 

579.  Aggregate  Fruits  are  those  in  which 
a  cluster  of  carpels,  all  belonging  to  one 
flower,  are  crowded  on  the  receptacle  into 
one  mass,  as  in  the  raspberry  and  black- 
berry taken  as  a  whole.  (Fig.  646.)    They 
may  be  aggregates  of  any  kind  of  simple 

fruits.  But  when  dry  and  not  coherent,  the  mass  would  simply 
and  properly  be  described  as  a  head  or  spike  of  carpels,  more 
commonly  of  akenes,  as  in  Ranunculus,  Ane- 
mone, &c.  Yet  when  numerous  carpels  thus 
compacted  become  fleshy,  and  sometimes  more 
or  less  coherent,  the  aggregate  may  need  to  be 
taken  into  account.  The  best  name  for  it  is 
that  of  SYNCAKPIUM,  or  in  English 
form  SYNCARP.  But  the  term  has 
been  applied  to  multiple  fruits  as 
well.1  In  Hydrastis,  the  numerous 
carpels  imbricated  on  the  upper 
part  of  the  torus  are  baccate,  that 
is,  become  berries  ;  in  a  raspberry, 
the  seemingly  baccate  grains  are  drupaceous  (being  drupelets,  573), 

1  The  syncarp  which  is  a  gynoecium  might  be  designated  a  simple  syn- 
carpium  ;  that  which  is  an  inflorescence,  a  complex  syncarpium,  which  may  be 
biflorous,  pauciflorous,  or  multiflorous. 

FIG.  645.  The  larger  Cranberry,  Vaccinium  (Oxycoccus)  macrocarpon ;  the  berry 
transversely  divided. 

FIG.  646.  Vertical  section  of  half  of  a  blackberry  (of  Rubus  villoeus),  enlarged ;  and, 
647,  of  one  of  its  drupelets  more  magnified. 


300 


THE   FRUIT. 


and,  slightly  cohering  together  (though  without  organic  union), 
they  fall  as  one  bod}'  from  the  conical  dry  torus  at  maturity.  It 
is  the  same  in  blackberries  or  bramble-berries  (Fig.  646,  647), 
except  that  the  drupelets  persist  on  the  torus,  which  partakes  of 
the  juiciness.1  In  the  aggregate  fruit  of  Magnolia  (Fig.  648-650), 
such  carpels,  imbricated  over  one  another,  cohere  more  or  less 

at  all  contiguous  parts,  and 
become  drupaceous ;  never- 
theless, at  maturity  each 
opens  dorsally,  allowing  the 
seeds  to  fall  out :  in  age  it 
dries  and  hardens,  and  also 
separates  from  its  connec- 
tions, and  so  be- 
comes a  follicle,  but 
with  the  remark- 
able peculiarity  of 
dorsal  instead  of 
ventral  dehiscence. 
(Fig.  650.)  In  Li- 
riodendron,  a  tree 
of  the  same  family, 
such  carpels  are 
dry  and  indehiscent  throughout ;  and  they  largely  consist  of  long 
and  flat  styles,  imbricated  in  a  cone,  but  separating  from  each 
other  and  from  the  slender  torus  at  maturity,  when  each  becomes 
a  samara. 

580.  Accessory  or  Anthocarpous  Fruits  are  those  of  which  some 
conspicuous  portion  of  the  fructification  neither  belongs  to  the 
pistil  nor  is  organically  united  with  it,  except  by  a  common 
insertion.  The  part  thus  imitating  a  fruit,  while  it  is  really  no 
part  of  the  pericarp,  is  sometimes  called  a  Pseudocarp,  or  an 
Anthocarp  or  Anthocarpium.  This  condition  may  occur  either 
in  simple,  in  aggregate,  or  in  multiple  fruits. 


1  The  aggregate  fruit  like  that  of  Rubus  (named  by  some  Conocarpium, 
by  others  an  sEterio,  Erythrostomum,  &c. )  was  termed  by  Dumortier  a  Drupe- 
turn.  A  similar  aggregation  of  baccate  carpels  he  termed  a  Baccetum ;  of 
follicles,  a  Fotticetum,  &c.  All  such  names  may  look  well  in  a  system ;  but 
they  are  both  superfluous  and  unmanageable  in  phytography. 

PIG.  648.  Aggregate  fruit  of  Umbrella-tree,  Magnolia  Umbrella,  reduced  In  size;  a 
seed,  from  a  lower  dehiscent  carpel  hangs  on  a  thread,  consisting  of  a  tuft  of  extensile 
spiral  ducts  unravelled.  649.  Same  in  longitudinal  section.  650.  One  of  the  carpels 
detached,  at  full  maturity,  dried  up,  dorsally  dehiscent,  exposing  the  pair  of  seeds  of 
the  natural  size. 


ITS  KINDS.  301 

581.  Gaultheria  procumbens,  the  aromatic  Wintergreen  (Fig. 
651,   652),  affords  a  good  example  of  the  first.     Its  seeming 
berry  (the  checkerberry) ,  with  summit  crowned  by  the  tips  of 
the  catyx-lobes,  well  imitates  the  true 

berry  of  a  Vaccinium,  such  as  that  of 
Fig.  645.  But  it  comes  from  a  flower 
with  thin  calyx,  underneath  and  free 
from  the  ovary.  Its  fruit  is  really 
a  capsule :  in  the  process  of  fructi- 
fication, the  calyx  enlarges,  becomes 
succulent,  completely  encloses  the  capsule  or  true  fruit,  yet 
without  adhering  to  it,  and  in  ripening  counterfeits  a  red 
berry.  So  in  Shepherdia,  or  Buffalo  Berry,  the  seeming  sarco- 
carp  of  a  drupe  is  really  a  free  calyx,  accrescent  and  succulent, 
enclosing  an  akene.  So,  also,  the  apparent  achenium  or  nut  of 
Mirabilis,  or  Four-o'clock,  and  of  its  allies, 
is  the  thickened  and  indurated  base  of  the 
tube  of  a  free  calyx,  which  contracts  at  the 
apex  and  encloses  the  true  pericarp  (a  utricle 
or  thin  akene) ,  but  does  not  cohere  with  it. 

582.  Likewise  the  torus,  although  not  con- 
spicuous, maybe  said  to  be  an  accessory  part 
of  the  aggregate  fruit  of  the  Blackberry  or 
Bramble  (579)  :   it  becomes  the  solely  con- 
spicuous and  the  sole  edible  part  of  a  straw- 
berry (389,  Fig.  406,  653),  the   akenes   or 
true  fruits  dispersed  over  the  surface  being 
apparently   insignificant.     Equally  in   many 

multiple  fruits  the  conspicuous  flesh  belongs  to  receptacle  (either 
torus  or  rhachis),  to  calyx,  or  even  in  part  to  bracts,  or  to  all 
these  parts  combined,  as  in  a  pine-apple. 

583.  Multiple  or  Collective  Fruits  *  are  those  which  result  from 
the  aggregation  of  several  flowers  into  one  mass.     The  simplest 
of  these  are  those  of  the  Partridge-Berry  (Mitchella,Fig.467), 


1  Collective  is  the  preferable  name.  The  term  multiple  was  applied  by 
DeCandolle  to  what  are  here  (following  Lindley)  called  aggregate  fruits ; 
and  the  aggregate  fruits  of  DeCandolle  are  here  called  multiple  or  collective. 
Moreover,  the  distinction  between  accessory  or  anthocarpous  and  collective 
or  multiple  fruits  was  not  recognized  by  Lindley,  who  combined  the  two 
in  his  original  "Introduction  to  Botany."  In  this  work  four  classes  are 
given :  1.  Fruit  simple,  APOCARPI  ;  2.  Fruit  aggregate,  AGGREGATI  ; 

FIG.  651.  Forming  capsule  of  Gaultheria  procumbens,  with  enlarging  calyx  partly 
covering  it.  652.  Same,  more  advanced,  and  in  longitudinal  section. 

FIG.  653.    Vertical  section  of  half  a  strawberry.    Compare  with  Fig.  406. 

LIBRARY 


STATE  TEACHER   COLUEOfc 
»WTA   BARBARA..   CAu«FQ«N1A 


SANTA   BAHUAR 


302 


THE   FRUIT. 


and  of  certain  species  of  Honeysuckle,  formed  of  the  ovaries 
of  two  blossoms  united  into  one  fleshy  fruit.  The  more  usual 
sorts  are  such  as  the  pine-apple,  mulberry,  and  the  fig.  These 
are,  in  fact,  dense  forms  of  inflorescence,  with  the  fruits  or  floral 
envelopes  matted  together  or  coherent  with  each  other ;  and  all 
or  some  of  the  parts  succulent.  The  grains  of  the  mulberry 
(Fig.  654-656)  are  not  the  ovaries  of  a  single  flower,  like  those 


of  the  blackberry,  which  it  superficially  resembles  :  the}'  belong 
to  as  many  separate  flowers  ;  and  the  pulp  pertains  to  the  calyx, 
not  to  the  pericarp,  which  is  an  akene.  So  that  this,  like  most 
multiple  fruits,  is  anthocarpous  as  well  as  multiple.  Similarly, 
the  mostly  indefinite  fructiferous  masses  of  Strawberry  Elite  may 
resemble  strawberries  ;  but  the  pulp}r  part  is  the  calyx  of  many 
flowers,  not  the  succulent  receptacle  of  one.  In  the  pine-apple, 
the  flowers  are  spicate  or  capitate  on  a  simple  axis,  which  grows 
on  beyond  them  into  leafy  stem ;  this  when  rooted  as  a  cutting 


3.  Fruit  compound  (ovaria  compound),  SYNCARPI  ;    4.  Collective  fruits, 
Anthocarpi. 

Later,  in  his  "Elements  of  Botany,"  Lindley  reduced  the  classes  to 
two:  1.  Simple  Fruits,  those  proceeding  from  a  single  flower;  2.  Multiple 
^fruits,  those  formed  out  of  several  flowers. 

FIG.  654.  A  mulberry,  youug.  655.  One  of  the  fleshy  grains  at  flowering  time,  show- 
Ing  It  to  be  a  pistillate  blossom  with  fleshy  calyx.  656.  The  same  later,  with  the  succu- 
lent sepals  in  transverse  section. 

PIG.  657.  A  young  fig.  658.  Longitudinal  section  of  the  same  later,  but  in  flowering 
time.  659.  A  small  slice,  magnified,  showing  some  of  the  flowers. 


ITS  KINDS.  303 

bears  another  pine-apple,  and  so  on :  the  constituent  flowers  have 
through  immemorial  propagation  in  this  way  become  sterile  and 
seedless,  and  all  its  parts,  along  with  the  bracts  and  the  axis  of 
the  stem,  blend  in  ripening  into  one  fleshy  and  juicy  mass.  Few 
fruits  of  this  class  have  ever  been  technically  named,  at  least 
with  names  which  have  come  into  use.  But  the  two  following 
deserve  special  appellations,  although  only  the  latter  is  familiar 
either  in  ordinary  language  or  in  descriptive  botany. 

584.  The  Syconium  or  Hypanthodium,  the  Fig  fruit.     (Fig.  657- 
659.)     This  results  from  a  multitude  of  flowers  concealed  in  a 
hollow  flower-stalk,  if  it  may  be  so  called,  which  becomes  pulpy 
and  edible  when  ripe  ;  and  thus  the  fruit  seems  to  grow  directly 
from  the  axil  of  a  leaf,  without  being  preceded  by  a  blossom. 
The  minute  flowers  within,  or  some  of  them,  ripen  their  ovaries 
into  very  small  akenes,  which  are  commonly  taken  for  seeds. 
The  fig  is  to  the  mulberry  what  a  rose-hip  is  to  a  strawberry. 
(389,  Fig.  406,  407.)     It  is  further  explained  by  a  comparison 
with  a  near  relative  of  the  Fig-tree,  Dorsteuia,  in  which  similar 
flowers  cover  the  upper  surface  of  a  flat  peltate  disk.     This  disk 
or  plate  sometimes  becomes  saucer-shaped  by  an  elevation  or 
incurvation  of  the  margin.      A  greater  degree  of  this  would 
render  it  cup- shaped,  or  even  pitcher-shaped  ;  from  which  it  is 
a  short  step  to  the  contraction  of  the  mouth  down  to  the  small 
orifice  which  is  found  in  the  fig. 

585.  The  Strobile  or  Cone  (Fig.  660)  is  a  scaly  multiple  fruit, 
resulting  from  the  ripening  of  certain  sorts  of  catkin.    The  name 
is  applied  to  the  fruit  of  the  Hop,  where 

the  large  and  thin  scales  are  bracts ; 
but  it  more  especially  belongs  to  the 
Pine  or  Fir  cone,  the  peculiar  fruit  of 
Coniferse  (507),  in  which  naked  seeds 
are  borne  on  the  upper  face  of  each 
fructiferous  scale  (Fig.  661),  or  some- 
times in  their  axils. 

Such  a   cone  when 

spherical,    and     of 

thickened        scales 

with  narrow  base,  as 

that  of  Cypresses, 

has  been  termed  a 
GALBULUS,  an  unnecessary  name.  The  galbulus  of  Juniper  is  a 

FIG.  660.  Strobile  or  Cone  of  a  Pitch  Pine,  Pinus  riglda.  661.  Inside  view  of  one 
of  the  scales,  showing  one  of  the  winged  seeds,  and  the  place  from  which  the  other,  662, 
has  been  detached. 


304  THE  FRUIT. 

remarkable  transformation  into  a  seeming  berry  ;  the  few  scales 
cohering  with  each  other  as  they  grow  and  becoming  fleshy  at 
maturity,  completely  enclosing  a  few  bony-coated  seeds. 

586.  A  Synopsis  of  the  kinds  of  Fruit,  as  characterized  in 
this  chapter,  is  appended.  The  analysis  extends  only  to  simple 
fruits.  For  there  are  no  commonly  used  special  names  of 
kinds  of  Aggregate  (579),  Accessory  (580),  or  Multiple  (583) 
fruits,  except  that  of  Strobile. 

SIMPLE  FRUITS  are 

Dry  and  dehiscent,  monocarpellary, 

Opening  by  one  (chiefly  the  ventral)  suture, FOLLICLE. 

Opening  by  both  sutures, LEGUME. 

Or  transversely  jointed, LOMENT. 

Dry  and  dehiscent,  bi-pluri-carpellary, CAPSULE. 

When  its  dehiscence  is  circumscissile, PYXIS. 

When  dehiscent  by  two  valves  from  two  parietal  placentae, .    .    SILIQUE. 

A  short  and  broad  silique, SILICLE. 

Dry  and  bi-pluri-carpellary,  splitting  into  one-seeded  carpels,    .  SCHIZOCARP. 

The  dimerous  schizocarp  of  Umbellif  erae, CREMOCARP. 

Each  of  its  halves  or  carpels, HEMICARP  or  MERICARP. 

The  akene-like  or  nut-like  parts  into  which  Schizocarps  generally 

divide, NUCULES  or  NUTLETS. 

Dry  and  indehiscent,  one-celled,  one-two-seeded, 

Winged, SAMARA. 

Wingless,  and  with  the 

Thin  pericarp  consolidated  with  the  seed, CARTOPSIS 

Thin  pericarp  loose  and  not  filled  by  the  seed, UTRICLE. 

Thick  or  hard  pericarp  free  from  the  seed, 

Small,  from  a  one-celled  one-two-ovuled  ovary,  AKENE  or  ACHENIUM. 
Larger,  mostly  from  a  two-several-celled  and  ovuled  ovary,  .       NUT. 

Nut  borne  in  a  cupule  or  involucre, GLANS. 

Fleshy  and  indehiscent, 

Heterogeneous  in  texture,  having 

A  stone  (putamen)  or  nutlets  within  an  exterior  sarcocarp,       .  DRUPE. 

Papery  or  cartilaginous  carpels  in  an  inferior  sarcocarp,     .    .     POME. 

A  harder  or  firm  rind  or  exterior,  and  soft  interior, 

From  an  inferior  ovary  (confined  to  Gourd  Family),     .     .    .      PEPO. 

From  a  superior  ovary  (confined  to  Orange  Family),     HESPERIDIUM. 

Homogeneous,  fleshy  throughout, BERRY. 


THE  SEED.  305 


CHAPTER  VIIL 

THE   SEED. 

587.  THE  SEED  is  the  fertilized  ovule   (515),  with  embryo 
formed  within  it.     It  consists,  like  the  ovule,  of  a  nucleus  or 
kernel,  enclosed  by  integuments.     The  seed-coats  are  those  of 
the  ovule,  viz.  two,  or  sometimes  only  one,  in  certain  plants 
none.     Occasionally  an  accessory  coat  appears  after  fertiliza- 
tion ;  and  certain  appendages  may  be  produced,  as  outgrowths 
from  some  part  of  its  surface  or  from  its  base.     The  nucleus  or 
kernel  is  composed  either  of  the  embryo  alone,  or  of  a  nutritive 
deposit  in  addition.    (19-41.)     All  the  parts  of  a  seed  are  in- 
dicated in  Fig.  663. 

588.  The  SEED-STALK  or  PODOSPERM,  when  there  is  one,  is 
the  funiculus  of  the  ovule  (516),  and  retains  this  name.     So 
also  do  the  CHALAZA,  RHAPHE,  and  HILUM  ;  the  latter  being  the 
scar  left  by  the  separation  of  the  seed  from 

its  funiculus  or  directly  from  the  placenta. 
The  foramen  of  the  ovule,  now  closed,  is 
the  MICROPYLE  of  the  seed. 

589.  The  terms  which  denote  the  char- 
acter of  the  ovule,  such  as  orthotropous, 

campylotropous,  amphitropous,  and  anatropous,  apply  equally  to 
the  resulting  seed. 

590.  Seed-Coats.     The  integuments  of  the  seed  answer  to  the 
primine  and  secundine  of  the  ovule.     The  main  seed-coat  is  the 
exterior  integument  of  the  ovule  when  there  is  more  than  one. 
Being  the  most  firm  coat,  and  not  rarely  crustaceous  in  texture, 
it  takes  the  name  of  TESTA,  which  is  equivalent  to  seed-shell. 
It  has  also  been  named  SPEKMODERM  (seed-skin)  ,  and  sometimes 
Episperm.     The  latter  name  (meaning  upon  the  seed)  is  best 
applied  to  the  pellicle  or  outer  layer,  sometimes  a  thick  one, 
which  the  testa  of  certain  seeds  forms.     The  testa  is  extremely 
various  in  form  and  texture,  is  either  close  and  conformed  to 


FIG.  663.  Vertical  magnified  section  of  the  (anatropous)  seed  of  the  American  Lin- 
den; with  the  parts  indicated,  viz.  the  hilum  (a);  testa  (6);  tegmen  (c);  albumen  (d); 
embryo  (e).  664.  Vertical  section  of  the  orthotropous  seed  of  Helianthemum  Cana- 
dense,  with  its  funiculus,  a. 


806 


THE   SEED. 


the  nucleus,  or  loose  and  cellular  (as  in  Pyrola-seeds) ,  or  vari- 
ously appendaged. 

591.  The  inner  coat,  called  TEGMEN  and  sometimes  ENDO- 
PLJEUKA,  when  present  is  always  conformed  to  the  nucleus,  and 
is  thin  or   soft  and   delicate.     Sometimes  it  is  inconspicuous 
through  cohesion  with  the  nucleus  or  with  the  inner  surface  of 
the  testa.     In  ovules  of  one  coat  it  is  necessarily  wanting. 

592.  Appendages  or  outgrowths  of  the  testa  generally  have 
reference  to   dissemination.     Two  characteristic  kinds  of  such 

appendages  are  the  wing  and  the  coma, 
both  pertaining  only  to  the  seeds  of  dehis- 
cent fruits  and  calculated,  by  rendering 
seeds  buoyant,  to  facilitate  dispersion  by 
the  wind.  The  wing  of  a  Pine-seed  (Fig. 
661,  662)  is  a  part  of  the  carpellary  scale 
upon  which  the  two  ovules  grew.  In 
Trumpet  Creeper  (Fig.  665),  an  entire 
wing  surrounds  the  body  of  the  seed. 
In  the  related  Catalpa  (Fig.  666),  it  is 
mainly  extended  from  the  two  ends,  and 
almost  dissolved  into  a  coma,  the  name 
given  to  the  tuft  of  soft  hairs  like  that 
which  forms  the  down  at  one  end  of  the 
seed  of  Milkweed  (Fig.  667),  and  of 
Epilobium,  and  at  both  ends  in  several 
Apocynaceae.  In  the  Cotton-plant,  very 
long  and  soft  hairs,  admirably  adapted  for 
spinning,  thickly  cover  the  whole  seed- 
coat.  The  wing  and  coma  of  seeds  are 
functional^  identical  with  the  wing  and  the  pappus  of  the  pericarp 
in  the  samara  and  the  akenes  of  Composite  (563,  564),  but 
morphologically  quite  unlike  them. 

593.  There  are  other  (mainly  microscopic)  structures  on  some 
seed-coats  which  come  usefully  into  play  in  arresting  farther 
dispersion  at  a  propitious  time  Or  place.     In  many  but  not  all 
Polemoniaceae   (notably  in  Collomia),  in  certain  Acanthaceae, 
such  as  Ruellia  tuberosa  (and  equally  in  certain  Composite  of 
the  Senecio  tribe  and  in  Salvias,  &c.,  among  Labiate,  where 
this  structure  is  transferred  to  akenes  and  nutlets) ,  the  testa  is 
coated  with  short  hairs,  which  when  wetted  burst  or  otherwise 
open  and  discharge  along  with  mucilage  one  or  more  very  atten- 

FIG.  665.    Winged  seed  ofTrnmpet  Creeper,  Tecoma  radicans.   666.  That  of  Catalpa, 
becoming  comose :  the  body  divided  lengthwise  through  the  embryo. 
FIG.  067.    Comose  seed  of  Milkweed,  ABclepias  Cornuti. 


ITS   COATS   AND   APPENDAGES. 


307 


uated  long  threads  (spiricles)  which  were  coiled  within.     These, 

protruding  in  all  directions  and  in  immense  numbers,  form  a 

limbus  of  considerable  size  around  the  seed,  and  evidently  must 

serve  a  useful  end  in  fixing  these  sma'l 

and  light  seeds  to  the  soil  in  time  of 

rain,  or  to  moist  ground,  favorable  to 

germination.     In  cress  and  flax-seed, 

the  abundant  mucilage  developed  when 

wetted  comes  from  the  gelatination  of 

epidermal    cell- walls,    and 

subserves  a  similar  use. 

594.  While  the  testa  in 
many  seeds  is  hard  and 
crustaceous  or  bony,  imitat- 
ing the  pericarp  of  a  nut,  in  others  (such  as  Pseonia)  it  becomes 
berry-like  (baccate) ,  and  in  Magnolia,  drupaceous.1  (Fig.  668- 
671.)  These  may  also  be  regarded  as  adaptations  for  dissemi- 


nation, here  by  the  agency  of  birds,  attracted  by  bright  coloring 
and  edible  pulp. 

595.  The  rhaphe  of  an  anatropous  seed  (shown  in  Fig.  681, 
685)  is  sometimes  so  salient  as  to  form  a  conspicuous  appen- 
dage, as  in  Sarracenia,  Fig.  672.  Again  it  may  be  wholly 

1  See  article  On  the  Structure  of  the  Ovule  and  Seed-coats  of  Magnolia, 
in  Jour.  Linn.  Soc.  ii.  106,  from  which  the  accompanying  figures  and  Fig. 
589-597  are  reproduced. 

FIG.  668.  Forming  seed  (one  eighth  of  an  Inch  long)  of  Magnolia  Umbrella;  the 
rhaphe  toward  the  eye.  669.  Magnified  view  of  the  same  divided  lengthwise  through 
the  rhaphe ;  the  outer  coat,  a,  beginning  to  form  a  hard  inner  layer,  af.  Within  and 
distinct  from  this  is  the  inner  coat  (b\  immediately  enclosing  the  nucleus,  c.  The  oppo- 
site side  of  the  testa  is  thicker  on  account  of  the  rhaphe,  in  which  d  indicates  the  cord 
of  spiral  ducts. 

FIG.  670.  A  nearly  full-grown  seed,  of  the  natural  size.  671.  Longitudinal  section, 
emargea,  showing  the  crustaceous  or  stony  inner  stratum  of  the  testa  well  developed: 
the  parts  lettered  as  in  Fig.  669.  672.  A  transverse  section  in  the  same  position. 


308  THE   SEED. 

inconspicuous,  as  in  the  ripe  seed  of  Magnolia,  where  it  is  at 
length  completely  merged  and  imbedded  in  the  fleshy  drupaceous 
testa,  as  shown  in  Fig.  670-672. 

596.  Crest- like  or  other  appendages  are  not  uncommon  either 
on  the  rhaphe  or  at  the  hilum.  These  are  outgrowths  produced 
during  the  development  of  the  ovule  into  the  seed.  In  Sangui- 
naria,  such  a  crest  develops  from  the  whole  length  of  the  rhaphe 


672 

(Fig.  673)  ;  in  Dicentra,  Corydalis  (Fig.  674),  &c.,  from  some 
part  of  it,  mostly  from  its  base  next  the  hilum,  or  from  the 
hilum  itself,  or  even  from  just  below  it.  Such  an  appendage, 
especially  when  attached  to  the  base  of  the  seed,  is  named  a 
STROPHIOLE.  A  similar  and  commonly  a  wart-shaped  appendage 
in  Euphorbia,  Ricinus  (Fig.  675),  dec.,  is  produced  by  an  out- 
growth of  the  external  orifice  of  the  ovule,  the  micropyle  of  the 
seed.  This  properly  takes  the  name  of  CARUNCLE.  But  the 
two  terms  are  not  always  discriminated.  B}r  further  develop- 
ment, either  of  these  ma3Tgive  rise,  in  certain  seeds,  to  an  acces- 
sory covering  called 

597.  The  Aril  or  Arillus.     This  term,  rather  vaguely  employed 
by  Linnaeus,  was  first  well  defined  by  Gaertner.     The  true  arillus 
is  an  accessory  seed-covering,  more  or  less  incom- 
plete, formed  between  the  time  of  fertilization  and 
the  ripening  of  the  seed,  by  a  growth  from  the  apex 
of  the  funiculus  (when  there  is  any)  at  or  just  be- 
low the  hilum,  in  a  manner  similar  to  that  in  which 
the  coat  or  coats  of  the  ovule  are   formed.     That 
876  of  Nymphaea  (Fig.  676)  is  a  typical  example  ;  only 

the  arillus  is  developed  from  the  funiculus  at  a  point  distinctly 
below  its  apex  :  here  a  ring  forms,  which  grows  into  a  cup,  and 
this  is  soon  extended  into  a  sac,  loosely  enclosing  the  seed,  and 
open  at  the  top.  This  is  membranaceous  ;  commonly  it  is  fleshy. 
When  there  is  absolutely  no  funiculus,  the  aril  may  originate 
from  the  placenta,  as  it  does  in  Podophyllum,  in  which  most  of 

FIG.  672.  Anatropous  seed  of  Sarracenia  purpurea,  with  very  salient  rhaphe.  673. 
Same  of  Sanguinaria  or  Bloodroot,  with  rhaphe  crested  for  its  whole  length.  674.  Seed 
of  Corydalis  aurea,  with  crest  or  strophiole,  attached  at  or  near  the  hilum.  675.  Seed 
(suspended)  of  Ricinus,  with  its  caruncle. 

FIG.  676.  Seed  of  White  Water-Lily,  Nymphsea  odorata,  in  its  loose  and  thin  arillufl. 


ABILLUS,   ALBUMEN.  309 

the  pulp  of  the  berry  consists  of  these  fleshy  arils,  much  com- 
pacted. (Fig.  677,  678.) 

598.  The  laciniate  aril  of  the  nutmeg  (mace)  and,  it  is  said, 
the  bright  red  and  pulpy  aril  of  Euonymus  and  Celastrus  begin 
in  the  manner  of  a  ca- 
runcle,  and   are  formed 

(mainly  if   not  wholly) 

of   an   outgrowth   at  or 

around  the  micropyle.  So 

that,  if  an  orthotropous 

seed   ever  developed  an 

aril  of  this  sort,  it  would 

be  seen  to  begin  at  the  apex  of  the  seed  and  cover  it  from  above 

downward.     Planchon,  who  distinguished  this  from  the  true  aril, 

gave  to  it  the  name  of  ARILLODE  (Arillodium)  or  False  Arillus. 

599.  The  Nucleus,  or  kernel  of  the  seed,  consists  of  the  Albu- 
men, when  this  substance  is  present,  and  the  Embryo. 

600.  The  Albumen,  as  described  in  the  second  chapter  (25,  &c.) , 
is  the  name  generally  employed  by  systematic  botanists  for  a 
store  of  nutritive  matter  in   the   seed  outside  of  the  embryo, 
whatever  its  chemical  composition.     It  is   not  here   the  name 
of  a  chemical  substance  (albumen  or  albumin) ,  but  of  a  cellular 
structure,  the  cells  of  which  are  loaded  commonly  with  starch- 
grains   (as  in  the  Cerealia),  more  or  less  mingled  with  other 
matters,  or  else  filled  with  an  encrusting  deposit  of  some  equiva- 
lent substance,  as  in  the  cocoanut,  coffee-grain,  &c.     The  cells 
in  which  this  deposit  is  made  belong  either  to  the  original  tissue 
of  the  nucleus,  or  to  a  new  formation  within  the  embryo-sac, 
mostly  to  the  latter.  (503.) 

601 .  Albumen  may  be  said  to  belong  to  all  seeds  in  the  grow- 
ing stage.     In  what  are  called  albuminous  seeds  it  persists  and 
forms  either  almost  the  whole  kernel,  the  embryo  remaining 
minute  (as  in  Fig.  23, 54,  680),  or  forms  a  large  portion  of  it  (Fig. 
13,  17,  19,  21,  48,  663,  664),  or,  by  the  growth  of  the  embryo 
displacing  it,  it  may  in  the  ripe  seed  be  reduced  to  a  thin  stratum 
or  mere  lining  to  the  contiguous  seed-coat ;  or  it  may  disappear 
altogether,  as  in  the  seeds  of  Maple,  Almond,  Squash,  Pea,  and 
the   like,    which  are  therefore  said   to  be  exalbuminous.     The 
difference  between  albuminous  and  exalbuminous  seeds  is  that 
the  maternal  nutritive  deposit  is  transferred  to  the  embryo  in 


FIG.  677.  Section  of  pericarp  and  placenta  of  Podophyllum  peltatum ;  the  pulp  of 
the  latter  mainly  of  the  nature  of  arillus,  investing  the  seeds.  678.  The  arillus  of  one 
seed  detached  and  enlarged,  divided  lengthwise,  showing  the  seed  within. 


310  THE   SEED. 

the  former  during  germination,  in  the  latter  during  the  growth 
of  the  seed. 

602.  The   albumen  was   named  Perisperm   by  Jussieu,  and 
Endosperm   by  Richard  (25,  note)  ;   but   neither  name  has  in 
sj-stematic  botany  displaced  the  earlier  one  of  Grew  and  Gart- 
ner.    But  both  names  have  recently  been  brought  into  use  to 
distinguish  between  two  kinds  of  albumen,  that  formed  within 
the  embryo-sac,  which  is  specifically  termed  ENDOSPERM,  and  that 
formed  without,  which  takes  the  name  of  PERISPERM.     This  use 
comports  with  the  etymology  of  the  two  words,  the  former  refer- 
ring to  a  comparatively  internal  and  the  latter  to  an  external 
portion  of  the  seed  or  kernel. 

603.  In  most  seeds  the  albumen  is  endosperm :  in  Canna  it 
is  all  perisperin.     In  Nymphaea  and  its  allies  (except  Nelum- 

bium,  which  has  none)  most  of  it  is  perisperm ; 
but  a  thin  and  condensed  layer  of  endosperm 
surrounds  the  embryo,  where  with  the  per- 
sistent embryo-sac  (or  the  apex  of  it)  it 
forms  the  fleshy  sac  in  which  the  embryo  is 
enclosed.  It  is  the  same  in  the  Pepper  Family 
(Fig.  679) ,  except  that  there  is  a  larger  quan- 
tity of  endosperm  or  inner  albumen. 

604.  When  the  nucleus  of  a  ripe  seed  is  hollow,  as  in  the 
cocoanut  and  nux  vomica,  the  formation  of  endosperm,  which 
usually  begins  next  the  wall  of  the  embryo-sac,  has  not  proceeded 
so  as  to  fill  the  cavity.     The  embryo- sac  in  the  cocoanut  attains 
enormous  size,  and  the  cavity  is  filled  by  the  milky  fluid. 

605.  The  texture  or  consistence  of  the  albumen  differs  greatly. 
It  is  farinaceous  or  mealy  when,  consisting  mainly  of  starch- 
grains,  it  may  readily  be  broken  down  into  a  powder, 
as  in  wheat,  buckwheat,  &c. ;  oily,  when  saturated 
with  a  fixed  oil,  as   in  poppy-seed ;  fleshy,  when 
more  compact,  but  readily  cut  with  a  knife,  as  in 
the  seed  of  Barberry ;  mucilaginous,  when  soft  and 
somewhat  pulpy,  as  in  Morning  Glory  and  Mallow, 
but  when  dry  it  becomes  fleshy  or  harder ;  corneous, 
when  of  the  texture  of  horn,  as  in  coffee  and  the 

seed  of  Caulophyllum  ;  and  even  bony,  as  in  the  vegetable  ivory, 
the  seed  of  Phytelephas.  It  is  mostly  uniform  ;  but  in  the  nutmeg, 


FIG.  679.  Longitudinal  magnified  section  of  a  seed  of  Black  Pepper ;  showing  the 
large  episperm,  the  small  endosperm  in  the  persistent  embryo-sac,  and  in  this  the 
minute  embryo. 

FIG.  680.  Longitudinal  section  of  a  seed  of  the  so-called  Papaw,  Aaimina  triloba, 
with  ruminated  albumen  and  minute  embryo. 


THE  EMBRYO?*  311 

in  the  seeds  of  Asimina  (Fig.  680)  and  all  the  Custard-Apple 
family,  it  is  marked  by  transverse  lines  or  divisions  (caused  by 
inflexions  or  growths  of  the  inner  seed-coat) ,  giving  a  section 
of  it  either  a  marbled  appearance,  or  as  if  it  had  been  slit  by 
incisions  :  it  is  then  said  to  be  ruminated. 

be  c 


606.  The  Embryo,1  being  an  initial  plantlet  or  individual  of  a 
new  generation,  is  of  course  the  most  important  part  of  the  seed. 
To  its  production,  protection,  and  support,  all  the  other  parts  of 
the  fruit  and  flower  are  subservient. 

607.  In  an  embrjx)  of  full  development,  namely,  one  in  which 
all  the  parts  are  manifest  antecedent  to  germination,  these  parts 
are  the  Caulicle,  otherwise  called  Radicle,  the   Cotyledons,  and 
the  Plumule.  (20,  30.)     The  first  is  the  initial 

axis  or  stem,  a  primary  internode ;  the  second 

consists  of  the  leaves  of  the  primary  node ;  the 

third  is  a  beginning  of  a  farther  growth  which 

is  to  develop  more  stem  and  leaves.     Such  an 

embryo  is  usually  unaccompanied  by  albumen, 

having  in  the  course  of  its  growth  taken  into 

itself  (mostly  into  the  cotyledons)  the  provision 

which  in  other  seeds  is  mainly  accumulated  external  to  it  until  it 

is  drawn  upon  in  germination. 


1  The  word  Embryo  or  Embryon  was  applied  to  this  body  in  plants  by 
Bonnet  (Considerations  sur  les  Corps  organisees),  in  1762,  and  was  introduced 
into  systematic  botany  at  about  the  same  time  (1763)  by  Adanson :  it  was 
taken  up  by  Gaertner  in  1788.  Jussieu  in  the  Genera  Plantarum  (1789)  held 
to  the  term  Corculum  (the  cor  seminis)  which  came  down  from  Caesalpinus. 

Being  the  germinal  part  of  the  seed,  the  embryo  of  the  plant,  like  that  of 
the  animal,  is  in  general  language  often  called  the  Germ. 

FIG.  681.  Seed  of  a  Violet  (anatropous),  enlarged;  with  hilum  or  scar  (a),  rhaphe 
(6),  and  chalaza  (c)  indicated.  682.  Vertical  section  of  the  same,  showing  the  straight 
embryo  in  the  axis  of  the  mealy  albumen. 

FIG.  683.  Vertical  section  of  the  (orthotropous)  seed  of  Buckwheat,  showing  the 
embryo  folded  round  in  the  mealy  albumen. 

FIG.  684.  Vertical  section  of  the  (anatropous)  seed  of  Elodea  Virginica,  the  embryo 
completely  filling  the  coats. 

FIG.  685.  Seed  of  Delphinium  tricorne  (anatropous),  enlarged;  the  hilum,  the 
rhaphe,  and  the  chalaza  lettered  as  in  Fig.  681.  686.  Vertical  section  of  the  same  with 
c,  the  chalaza,  d,  the  testa,  e,  the  tegmen,/,  the  albumen,  g,  the  minute  embryo  near 
the  hilum  (a). 

FIG.  687.  Embryo  of  the  Pumpkin,  with  its  short  radicle  and  large  and  flat  cotyl* 
dons,  seen  flatwise.  688.  A  vertical  section  of  the  same,  viewed  edgewise. 


312  THE   SEED. 

608.  The  opposite  extreme  is  an  embryo  (as  in  Fig.  686) 
which  appears  as  a  mere  speck  in  the  albumen,  but  in  which 
close  microscopical  inspection  may  commonly  reveal  some  differ- 
entiation, such  as  a  slight  notch  at  one  end   (that  farthest  re- 
moved from  the  micropyle)  of  a  dicotyledonous  embryo,  indicating 
the  future  cotyledons.     Indeed,  in  Monotropeae,  Orobanchacese, 
and  some  other  parasitic  dicotyledonous  plants,  and  in  Orchids 
among  the  monocotyledonous,  the  embryo  is  a  globular  or  oblong 
iparticle,  with  no  adumbration  of  organs  whatever  antecedent  to 
germination.     There  are  all  grades  between  the  most  rudimen- 
tary and  the  most  developed  embryos. 

609.  Under  the  circumstances   of  its  formation   (532),  the 
radicular  end  of  the  embryo  is  always  near  to  and  points  towards 
the  micropyle  of  the  seed,  viz.  to  what  was  the  orifice  of  the 
ovule  ;  and  if  the  embryo  be  straight,  or  merely  partakes  of  the 
curvature   of  the  seed,   the  cotyledons   point  to  the   opposite 
extremity,  that  is,  to  the  chalaza. 

610.  The  position  of  the  radicle  as  respects  the  hilum  varies 
with  the  different  kinds  of  seed.    In  the  orthotropous  form,  as  in 
Helianthemum  (Fig.  664)  and  Pepper  (Fig.  679),  the  radicle 
necessarily  points  directly  away  from  the  hilum.1     In  the  anatro- 
pous  form,  as  in  Fig.  663,  682,  and  684-686,  the  extremity 
of  the  radicle  is  brought  to  the  immediate  vicinity  of  the  hilum ; 

and  so  it  is,  although  in  a  different  way, 
in  the   campylotropous   seed   (Fig.  689, 
690) ;    while    in    the    amphitropous   the 
radicle  points  away  from  the  hilum  later- 
ess  ego  ally.    As  the  nature  of  the  ovule  and  seed 
may  usually  be  ascertained  by  external  inspection,  so  the  situation 

1  Two  technical  terms,  early  introduced  by  Richard  to  indicate  the  direc- 
tion of  the  radicle  (caulicle),  or  rather  its  relation  to  the  hilum,  are 

Antitropous,  when  the  embryo  directs  its  radicle  away  from  the  hilum,  as 
it  must  in  all  orthotropous  seeds ; 

Orthotropous,  also  homotropous,  when  directed  to  the  hilum  (more  strictly  to 
the  micropyle  close  to  the  hilum),  as  in  anatropous  seeds.  These  two  terms 
are  still  employed  by  many  botanists,  although  superfluous  when  the  ovule 
or  seed  is  stated  to  be  anatropous  or  orthotropous,  &c.  And  the  term 
orthotropous,  so  used,  is  liable  to  be  confused  with  orthotropous  as  applied 
to  the  ovule. 

Richard,  moreover,  termed  the  embryo  amphitropous  when  curved  or  coiled, 
as  in  Chickweed  (Fig.  689)  and  all  such  campylotropous  seeds ;  and  hetero- 
tropous  when  neither  radicle  nor  cotyledons  point  to  the  hilum,  as  occurs 
in  the  semi-anatropous  or  amphitropous  ovule.  Many  botanists  describe 
the  last  by  the  expression  "  radicle  vague,"  or,  better,  "  embryo  transverse." 

FIG.  689.  Campylotropous  seed  of  common  Chickweed,  magnified-  690.  Section  ol 
the  same,  showing  the  embryo  coiled  into  a  ring  around  the  albumen. 


THE  EMBRYO.  313 

of  the  embryo  within,  and  of  its  parts,  may  often  be  inferred 
without  dissection.  But  the  dissection  of  seeds  is  not  generally 
difficult. 

611.  The  direction  of  the  radicle  with  respect  to  the  pericarp 
is  also  noticed  by  systematic  writers ;  who  employ  the  terms 
radicle  superior  or  ascending  when  this  points  to  the  apex  of  the 
fruit ;  radicle  inferior  or  descending  when  it  points  to  its  base  ; 
centripetal,  when  turned  toward  the  axis  of  the  fruit ;  centrifugal 
(or  peritropous) ,  when  turned  toward   the  sides ;   and   vague, 
when  it  bears  no  evident  or  uniform  relation  of  the  kind  to  the 
pericarp. 

612.  The  position  of  the  embryo  as  respects  the  albumen, 
when  that  is  present,  is  various.     Although  more  commonly  in 
the  axis,  it  is  often  excentric,  or  even  external  to  the  albumen, 
as  in  all  Grasses  and  cereal  Grains  (Fig.  56-61),  inPolygonum, 
&c.     When  external  or  nearly  so,  and  curved  circularly  around 
the  albumen,  as  in  Chickweed  (Fig.  690)  and  Mirabilis  (Fig. 
17),  it  is  said  to  be  peripheric. 

613.  The  embryo  may  be  very  variously  folded  or  coiled  in 
the  seed.     The  two  cotyledons,  instead  of  plane  and  straight, 
may  be  crumpled  ;  or  they  may  be  simply  convolute  or  rolled  up 
from  one  edge,  as  in  Calycanthus  (Fig.  691)  ;  or  circinately  con- 


volute from  the  apex,  as  in  Bunias  ;  or  else  doubled  up  and  thus 
biplicately  convolute,  as  in  Sugar  Maple,  Fig.  2.  Two  modi- 
fications are  more  common,  and  are  of  such  classificatory  impor- 
tance in  Cruciferae  as  to  need  special  reference.  Namely,  when 
cotyledons  are 

Incumbent  (as  in  Fig.  692,  693),  being  so  folded  that  the  back 
of  one  is  laid  against  the  side  of  the  radicle  ;  and 

Accumbent  (Fig.  694,  695),  when  the  edges  of  the  pair  of 
cotyledons  are  longitudinally  applied  to  the  radicle.  These 
differences  were  first  employed  in  the  classification  Cruciferae  by 


FIG.  691.    Convolute  embryo  of  Calycanthus,  the  upper  half  cut  away. 

FIG.  692.  Seed  of  a  Cruciferous  plant  (Sisymbrium),  with  incumbent  cotyledons, 
divided.  693.  Embryo  of  the  same  detached  entire. 

FIG.  694.  Seed  of  a  Cruciferous  plant  (Barbarea)  with  accumbent  cotyledons. 
695.  The  embryo  entire. 


314  THE  SEED. 

Robert  Brown,  and  were  adopted  as  primary  and  tribal  characters 
by  DeCandolle. 

614.  As  to  number  of  cotyledons,  the  two  types  of  embryo 
are  the 

Monocotyledonous,  with  a  single  cotyledon,  i.  e.  leaves  at  the 
first  nodes  alternate  (39)  ;  and 

Dicotyledonous,  with  a  pair  of  cotyledons,  i.  e.  leaves  of  the 
first  node  in  the  most  simple  whorl,  a  pair,  in  other  words,  oppo 
site  (21)  ;  with  its  modification  of 

Polycotyledonous  (38),  the  leaves  of  the  first  node  in  whorls  of 
three,  four,  or  more.  This  occurs  with  constancy  in  a  majority 
of  Coniferae  (Fig.  48,  49) ,  occasionally  and  abnormally  in  sundry 
ordinary  dicotyledonous  species. 

615.  There  are  several  embryos  of  the  cotyledonous  type  in 
which  one  cotyledon  is  smaller  than  the  other,  viz.  the  inner 
one  when  the  emb^o  is  coiled  or  folded.     And  in  all  the  species 
of  Abronia  (a  genus  allied  to  Mirabilis,  Fig.  18)  this  cotyledon 
is  wanting,  so  that  the  embryo  becomes  technically  monocotyle- 
donous.     In  another  genus,  the  Dodder  (Fig.  78,  79),  both 
cotyledons  are  constantly  wanting  ;  and  the  plumule  shows  only 
minute  scales,  the  homologues  of  succeeding   leaves    reduced 
almost  to  nothing. 

616.  Sometimes  the  two  cotyledons  are  consolidated  into  one 
body  by  the  coalescence  of  their  contiguous  faces ;  when  they 
are  said  to  be  conferruminate.     This  occurs  more  or  less  in  the 
Horsechestnut  and  Buckeye  (Fig.  41,  42),  and  is  striking  in 
the  seed  of  the  Live  Oak,  Quercus  virens, 

617.  The  general  morphology  of  the  embryo  and  its  develop- 
ment in  germination  were  described  at  the  commencement  of  this 
volume.    And  so  the  completion  of  this  account  of  plant,  flower, 
fruit,  seed,  and  embryo  brings  the  history  round  to  the  starting 
point.    (12-19,    &c.)      Having  mastered  the  morphology  and 
general  structure  of  the  higher  grade  of  plants,  the  pupil  may 
go  on  to  the  morphology  and  structure  of  cells  (or  Vegetable 
Anatomy  or  Histology),  and  to  the  study  of  Cryptogamous 
Plants  in  all  their  grades. 


TAXONOMY.  315 


CHAPTER  IX. 

TAXONOMY. 

SECTION  I.    THE  PRINCIPLES  OF  CLASSIFICATION  IN  NATURAL 
HISTORY. 

618.  TAXONOMY,  from  two  Greek  words  which  signify  arrange- 
ment and  law,  is  the  study  of  classification.     This  is  of  utmost 
importance  in  Natural  History,  on  account  of  the  vast  number 
of  kinds  to  be  set  in  order,  and  of  relations  (of  agreement  and 
difference)  to  be  noted.     Botanical  classification,  when  complete 
and  correct,  will  be  an  epitome  of  our  knowledge  of  plants. 
Arrangement  according  to   kinds,  and  of  special  kinds   under 
the  more  general,  is  common  to  all  subjects  of  study.     But  the 
classification  in  Biological  Natural  History,  that  is  in  Botany  and 
Zoology,  has  a  foundation  of  its  own. 

619.  The  peculiarity  of  plants  and  animals  is  that  they  exist 
as  individuals,  propagating  their  like  from  generation  to  genera- 
tion in  a  series.     Of  such  series  of  individuals  there  are  very 
many  kinds,  and  the  kinds  have  extremely  various  and  unequal 
degrees  of  resemblance.     There  are  various  gradations,  but  not 
all  gradations  of  resemblance.     Between  some,  the  difference  is 
so  wide  that  it  can  be  said  only  that  they  belong  to  the  same 
kingdom ;  between  others,  the  resemblance  is  so  close  that  it 
may  be  questioned  whether  or  not  they  came  from  common 
parents  or  near  ancestors. 

620.  The  recognition  of  the  perennial  succession  of  similar 
individuals  gives  the  idea  of  SPECIES.     The  recognition  of  un- 
equal degrees  of  likeness  among  the  species  is  the  foundation 
of  GENERA,  ORDERS,  CLASSES,  and  other  groups  of  species. 

621.  Individuals  are  the  units  of  the  series  which  constitute 
species.     The  idea  of  individuality  which  we  recognize  through- 
out the  animal  and  vegetable  kingdoms  is  derived  from  ourselves, 
conscious   individuals,   and   from  our  corporeal   structure  and 
that  of  the  higher  brute  animals.     This  structure  is  a  whole, 
from  which  no  part  can  be  abstracted  without  mutilation.     Each 
individual  is  an  independent  organism,  of  which  the  component 
parts  are  reciprocally  means  and  ends.     Individuality  is  a  main 


316  TAXONOMY. 

distinction  between  beings  and  things  ;  but,  although  the  tend- 
ency to  individuation  begins  with  life  itself,  it  is  completely 
realized  only  in  the  higher  animals. 

622.  In  plants,  as  also  in  some  of  the  lower  animals,  individu- 
ality is  merged  in  community.     No  plant  (except  one  reduced 
to  the  simplicity  of  a  single  cell,  of  circumscribed  growth,  and 
without  organs)  is  an  individual  in  the  sense  that  a  man  or  a 
dog  is.    (16,   156.)      The   herb,   shrub,   and   tree  are   neither 
indivisible  nor  of  definite  limitation.     Whether  their  successive 
growths  are  to  remain  parts  of  the  previous  plant,  or  to  be  inde- 
pendent plants,  depends  upon  circumstances ;  and  there  is  no 
known  limit  to  budding  propagation. 

623.  There  is,  however,  a  kind  of  social  or  corporate  indi- 
viduality in  those  animals,  or  communities   (whichever  we  call 
them)  of  the  lower  grade  which  are  multiplied  by  buds  or  off- 
shoots as  well  as  by  ova,  and  in  which  the  offspring  remains,  or 
may  remain,  organically  connected  with  the  stock.     The  poly- 
pidom  or  polyparium  commonly  has  a  certain  limitation  and  a 
definite  form ;   and   certain   polyps   may  become   organs   with 
special  functions  subordinate  to  the  common  weal.      This  is 
more  largely  true  in  the  vegetable  kingdom.     So  that  for  de- 
scriptive purposes,  and  in  a  just  although  somewhat  loose  sense, 
the  herb,  shrub,  or  tree  is  taken  as  an  individual.     But  only 
while  it  forms  one  connected  body.     Offshoots  when  separately 
established  are  equally  individuals  in  this  sense. 

624.  What  it  is  in  plants  which  philosophically  answers  to  the 
individual  in  the  higher  animals  is  another  question,  to  which 
various  answers  have  been  given.1     Some  insist  that  the  whole 
vegetative  product  of  one  seed  makes  one  individual,  whether 
connected  or  separated  (as  may  happen)  into  a  million  of  plants. 
But  a  common  and  less  strained  view  restricts  the  individual 
to  such  product  only  while   organically   united.      Others    (of 
which  Thouars  at  the  beginning  and  Braun  at  the  middle  of  the 
present  century  are  leading  examples)  take  each  axis  or  shoot 
with  its  foliage  to  represent  the  individual,  of  which  the  leaves 
and  their  homologues  are  organs,  the   branches  being  usually 
implanted  upon  the  parent  axis  as  this  is  implanted  in  the  soil, 
but  also  equally  capable  of  producing  roots  by  which  they  may 
make  their  own  connection  with  the  soil.     Still  others,  on  pre- 


1  For  the  history  of  opinion  upon  and  a  full  presentation  of  this  topic, 
see  Alexander  Braun's  Memoir  (originally  published  in  the  Abhandl.  Akad. 
Wissenschaften  zu  Berlin,  1863),  Das  Individuum  der  Pflanze,  &c.,  and  a 
translation  by  C.  F.  Stone  in  Amer.  Jour.  Sci.  ser.  2,  xix.  xx.  1855. 


THE  PRINCIPLES   OF   CLASSIFICATION.  317 

cisely  similar  grounds,  carry  the  analysis  a  step  farther,  and 
regard  each  phytomer  (16)  as  the  individual.  Finally,  some,  in 
view  of  their  potentially  independent  life,  take  the  cells,  or  units 
of  anatomical  structure,  to  be  the  true  individuals ;  and  this 
with  sufficient  reason  as  regards  the  simplest  cryptogamous 
plants.  Upon  the  view  here  adopted,  that  plants  do  not  rise 
high  enough  in  the  scale  of  being  to  reach  true  individuality, 
the  question  is  not  whether  it  is  the  cell,  the  phytomer,  the  shoot, 
the  tree,  or  the  whole  vegetative  product  of  a  seed  which  answers 
to  the  animal  individual,  but  only  which  is  most  analogous  to  it. 
In  our  view,  its  analogue  is  the  cell  in  the  lowest  grades  of  vege- 
table life,  the  phytomer  in  the  higher.1  But,  in  botanical  de- 
scription and  classification,  by  the  individual  is  meant  the  herb, 
shrub,  or  tree,  unless  otherwise  specified. 

625.  Species  in  biological  natural  history  is  a  chain  or  series 
of  organisms  of  which  the  links  or  component  individuals  are 
parent  and  offspring.     Objectively,  a  species  is  the  totality  ot 
beings  which  have  come  from  one  stock,  in  virtue  of  that  moU 
general  fact  that  likeness  is  transmitted  from  parent  to  progenj-. 
Among  the  many  definitions,  that  of  A.  L.  Jussieu  is  one  of  the 
briefest  and  best,  since  it  expresses  the  fundamental  conception 
of  a  species,  *.  e.  the  perennial  succession  of  similar  individuals 
perpetuated  by  generation. 

626.  The  two  elements  of  species  are  :  1,  community  of  origin  ; 
and,  2,  similarity  of  the  component  individuals.     But  the  degree 
of  similarity  is  variable,  and  the  fact  of  genetic  relationship  can 
seldom  be  established  by  observation  or  historical  evidence.     It 
is  from  the  likeness  that  the  naturalist  ordinarily  decides  that 
such  and  such  individuals  belong  to  one  species.     Still  the  like- 
ness is  a  consequence  of  the  genetic  relationship ;  so  that  the 
latter  is  the  real  foundation  of  species. 


1  For  just  as  successive  branches  are  repetitions  and  progeny  of  the 
parent  branch  or  stem,  the  phytomers  of  the  branch  are  repetitions  and 
progeny  each  of  the  preceding  one,  so  forming  a  series  of  vegetative 
generations ;  and  the  whole  tree  might  almost  as  well  represent  the  individ- 
ual as  one  of  its  branches.  The  phytomer,  as  well  as  the  branch,  is  capable 
of  completing  itself  by  producing  roots,  but  is  itself  indivisible  except  by 
mutilation.  Least  tenable  of  all  is  the  conception  that  the  whole  product 
of  a  seed  may  be  taken  to  represent  the  vegetable  individual.  For  then 
individuals  increased  by  buds  and  division  are  wholly  unlimited  both  in  ex- 
tent and  in  duration,  so  far  as  observation  can  show,  and  a  multitudinous 
race,  not  only  of  the  present  and  past,  but  perhaps  in  perpetuity,  may  con- 
sist of  a  single  individual.  There  are,  indeed,  theoretical  reasons  for  infer- 
ring that  a  bud-propagated  race  may  not  last  so  long  as  a  seed-propagated 
specie* :  but  there  is  no  proof  of  it.  See  Darwiniana,  Art.  xii. 


318  TAXONOMY. 

627.  No  two  individuals  are  exactly  alike ;  and  offspring  of 
the  same  stock  ma}'  differ  (or  in  their  progeny  may  come  to  differ) 
strikingly  in  some  particulars.     So  two  or  more  forms  which 
would  have   been  regarded   as  wholly  distinct   are  sometimes 
proved  to  be  of  one  species  by  evidence  of  their  common  origin, 
or  more  commonly  are  inferred  to  be  so  from  the  observation  of 
a  series  of  intermediate  forms  which  bridge  over  the  differences. 
Only  observation  can  inform  us  how  much  difference  is  compat- 
ible with  a  common  origin.     The  general  result  of  observation 
is  that  plants  and  animals  breed  true  from  generation  to  genera- 
tion within  certain  somewhat  indeterminate  limits  of  variation ; 
that  those  individuals  which  resemble  each  other  within  such 
limits  interbreed  freely,  while  those  with  wider  differences  do 
not.     Hence,  on  the  one  hand,  the  naturalist  recognizes  Varieties 
or  differences  within  the  species,  and  on  the  other  Genera  and 
other  superior  associations,  indicative  of  remoter  relationship  of 
the  species  themselves. 

628.  Varieties  are  forms  of  species  marked  by  characters  of 
less  fixity  or  importance  than  are  the  species  themselves.     They 
may  be  of  all  grades  of  difference  from  the  slightest  to  the  most 
notable :    they  abound  in   free   nature,  but  assume  particular 
importance  under  domestication  and  cultivation ;  under  which 
variations  are  most  prone  to  originate,  and  desirable  ones  are 
preserved,  led  on  to  further  development,  and  relatively  fixed. 

629.  If  two  seeds  from  the  same  pod  are  sown  in  different 
soils,  and  submitted  to  different  conditions  as  respects  heat,  light, 
and  moisture,  the  plants  that  spring  from  them  will  show  marks 
of  this  different  treatment  in  their  appearance.     Such  differences 
are  continually  arising  in  the  natural  course  of  things,  and  to 
produce  and  increase  them  artificially  is  one  of  the  objects  of 
cultivation.     Striking  as  they  often  are  (especially  in  annuals 
and  biennials) ,  they  are  of  small  scientific  consequence.     When 
spontaneous  they  are  transient,  the  plant  either  outlasting  the 
modifying  cause  or  else  succumbing  to  its  continued  and  graver 
operation.     But,  in  the  more  marked  varieties  which  alone  de- 
serve the   name,  the  cause  is   occult   and   constitutional ;   the 
deviation  occurs  we  know  not  why,  and  continues  throughout 
the  existence  and  growth  of  the  herb,  shrub,  or  tree,  and  con- 
sequently through  all  that  proceeds  from  it  by  propagation  from 
buds,  as  by  offsets,  layers,  cuttings,  grafts,  &c. 

630.  Some  varieties  of  cultivation  originate  in  comparatively 
slight  deviations  from  the  type,  and  are  led  on  to  greater  differ- 
ences  by  strict   selection   of  the  most  marked  individuals  to 
breed  from.     Most  appear  as  it  were  full-fledged,  except  as  to 


THE   PRINCIPLES   OF   CLASSIFICATION.  319 

luxuriance  or  development,  more  or  less  under  the  control  of 
conditions,  their  origin  being  wholly  unaccountable.  They  arise 
in  the  seed-bed,  or  sometimes  from  buds,  which  as  the  gardeners 
say  "  sport."  x  That  is,  some  seedlings,  or  some  shoots,  are 
unlike  the  rest  in  certain  particulars.2 

631.  Most  varieties  originate  in  the  seed,  and  therefore  the 
foundation  for  them,  whatever  it  may  be,  is  laid  in  sexual  repro- 
duction.    But  Bud-variation,  or  the  "  sporting"  of  certain  buds 
into  characters  in  branch,  flower,  or  fruit  unlike  those  of  the 
stock,  is  known  in  a  good  number  of  plants.8     It  might  also 
occur  in  corals,  hydras,  and  other  compound  animals  propagated 
by  budding.     Once   originated,  these  varieties  mostly  persist, 
like  seedling  varieties,  through  all  the  generations  of  budding 
growth,  but  are  not  transmitted  to  the  seed. 

632.  Upon  the  general  principle  that  progeny  inherits  or  tends 
to  inherit  the  whole  character  of  the  parent,  all  varieties  must 
have  a  tendency  to  be  reproduced  by  seed.     But  the  inheritance 
of  the  new  features  of  the  immediate  parent  will  commonly  be 
overborne  by  atavism,  i.  e.  the  tendency  to  inherit  from  grand- 
parents, great-grand-parents,  &c.  Atavism,  acting  through  a  long 
line  of  ancestry,  is  generally  more  powerful  than  the  heredity 
of  a  single  generation.     But  when  the  offspring  does  inherit  the 
peculiarities  of  the  immediate  parent,  or  a  part  of  them,  its  off- 
spring has  a  redoubled  tendency  to  do  the  same,  and  the  next 
generation  still  more  ;  for  the  tendency  to  be  like  parent,  grand- 
parent, and  great-grand-parent  now  all  conspire  to  this  result 
and  overpower  the  influence  of  remoter  ancestry.     Close-breed- 
ing (398)  is  requisite  to  this  result.     In  the  natural  wild  state, 
varieties  —  many  and  conspicuous  as  they  often  are  —  must  be 
much  repressed  by  the  prevalent  cross-fertilization  which  takes 
place  among  the  individuals  of  almost  all  species.    Cultivators  and 
breeders  in  fixing  varieties  are  careful  to  secure  close  breeding 
as  far  as  this  is  possible.     This  has  fixed  the  particular  sorts  of 
Indian  Corn,  Rye,  Cabbage,  Lettuce,  Radishes,  Peas,  &c.,  and 

1  Both  the  technical  English  term,  Sport,  and  its  Latin  equivalent,  Lusus, 
are  sometimes  used  for  bud-variation  only,  yet  as  commonly  for  seedling 
variation  also. 

2  Darwin  assumes  that  variation  is  of  itself  indefinite  or  vague,  tending 
in  no  particular  direction,  but  that  direction  is  wholly  given  by  the  elimina- 
tion in  the  struggle  for  life  of  all  but  the  fittest  for  the  conditions.     But 
what  we  observe  in  the  seed-bed  does  not  suggest  this  view.     Naegeli,  Braun, 
and  myself  incline  to  the  opinion  that  each  plant  has  an  inherent  tendency 
to  variation  in  certain  general  directions. 

8  A  list  of  known  bud-varieties  is  given  in  Darwin's  Variation  of  Animals 
and  Plants  under  Domestication,  Chapter  xi. 


320  TAXONOMY. 

indeed  of  nearly  all  our  varieties  of  cultivated  annual  and  biennial 
esculent  plants,  as  well  as  of  several  perennials,  many  of  which 
have  been  fixed  through  centuries  of  domestication,  while  others 
are  of  recent  establishment.  What  is  now  taking  place  with 
the  Peach  in  this  country  may  convince  us  that  heritable  varieties 
may  be  developed  in  trees  as  well  as  in  herbs,  and  in  the  same 
manner ;  and  that  the  reason  why  most  races  are  annuals  or 
biennials  is  because  these  can  be  perpetuated  in  no  other  way, 
and  because  the  desired  result  is  obtainable  in  fewer  years  than 
in  shrubs  or  trees.  Varieties  of  this  fixity  of  character  are  called 

633.  Races  (Lat.  Proles).     A  race,  in  this  technical  sense  of 
the  term,  is  a  variety  which  is  perpetuated  with  considerable 
certainty  by  sexual  propagation.     This  distinction  of  varieties 
pertains  chiefly  to  botany.     In  the  animal  kingdom  all  permanent 
varieties  must  be  races.      So  are  all  indigenous  varieties  of 
plants.1     In  most  of  these,  the  position  of  species  and  variety  is 
more  or  less  arbitrary  or  accidental,  and  capable  of  interchange. 
What  is  called  the  species  may  be  only  a  commoner  or  better- 
known  form,  or  the  one  first  recognized  and  named  by  botanists  ; 
whence  the  other  forms  as  they  come  to  be  recognized  are  made 
to  rank  in  the  books  as  varieties.     Instead  of  one  varying  from 
the  other,  all  the  forms  have  probably  varied  ages  ago  from  a 
common  type. 

634.  These  varieties  of  the  highest  order  and  most  marked 
characteristics,  being  perpetuable  by  seed,  have  the   principal 
attributes  of  species.     They  are  a  kind  of  subordinate  derivative 
species.      Hence   they   are   sometimes   called  Subspecies.     We 
judge  them  not  to  be  so  many  species,  either  because  in  the  case 
of  cultivated  races  we  know  something  of  their  origin  or  history, 
and  more  of  the  grave  changes  which  long  domestication  may 
bring  to  pass;   or  because   the   forms,  however  stable,  differ 
among  themselves  less  than  recognized  species  generally  do  ;  or 
because  very  striking  differences  in  the  extremes  are  connected 
by  intermediate  forms.     And  our  conclusions,  it  must  be  under- 
'stood,  "are  not  facts,  but  judgments,  and  largely  fallible  judg- 
ments." 2    For  while  some  varieties  appear  strikingly  different, 
some  species  are  very  much  alike.8 

1  The  Horseradish  and  a  few  other  plants  of  spontaneous  growth,  which 
through  long  dependence  on  bud-propagation  seem  to  have  lost  the  power 
of  setting  seed,  can  hardly  be  called  varieties. 

2  Darwiniana,  35. 

8  Wherefore,  since  we  hardly  need  the  term  race  in  the  restricted  sense 
of  seed-propagated  variety,  it  is  sometimes  convenient  to  use  it  in  the  man- 
ner proposed  by  Bentham  (Anniversary  Address  to  the  Linnean  Society, 


THE  PRINCIPLES   OF  CLASSIFICATION.  321 

635.  One  distinction  between  varieties  and  species  is  note- 
worthy and  important,  even  if  it  may  not  serve  as  a  criterion. 
The  individuals  of  different  varieties  in  plants  interbreed  as  freely 
as  do  those  of  the  same  variety  and  are  equally  prolific.     Their 
union  produces 

636.  Cross-breeds.1     In  nature,  cross-breeding  doubtless  re- 
presses variation  or  prevents  the  segregation  of  varieties  into 
what  would  be  ranked  as  species.     In  cultivation  and  domesti- 
cation, it  is  turned  to  important  account  in  producing  intermediate 
new  varieties  (cross-breeds)  variously  combining  the  different 
excellencies  of  two  parent  individuals  or  two  varieties.     Thus 
the  great  number  of  forms  produced  by  variation  (especially  as 
to  flowers  and  fruits)  have  been  further  diversified,  and  selected 
forms  improved  for  special  uses  by  judicious  combination. 

637.  In  general,  the   individuals  of  distinct  species  do  not 
interbreed,   although  many  are  capable  of  it.     There  is  great 
diversity  in  this  regard  among  plants,  some  (such  as  Willows, 
Verbascums,  and  Verbenas)  interbreeding  freely  and  reciprocally  ; 
some  interbreeding  in  one  direction,  but  not  reciprocally  ;  others, 
even  when  very  similar,  refusing  to  unite.     But,  on  the  whole, 
there  seems  to  be  few  nearly  related  species  in  which  the  pollen 
of  the  one  cannot  be  made  to  act  upon  the  ovules  of  the  other 
by  persistent  and  proper  management.     Such  crossing  is  an 
important  resource  in  horticulture.     Crossing  of  species,  when 
successful,  produces 

638.  Hybrids.    In  these,  the  characteristics  of  the  two  species 
are  combined,  sometimes  in  equal  proportions,  sometimes  with 
great  preponderance  of  one  or  the  other  parent ;  and  there  is 
often  a  difference  in  the  result  in  reciprocal  fertilizations.     Hy- 
brids do  not  play  a  very  prominent  part  in  nature,  apart  from 
cultivation,  although  the  limits  of  some  species  may  be  obscured 
by  them,  possibly  of  more  than  is  generally  supposed.     In  the 
animal  kingdom,  all  the  most  familiar  hybrids  are  sterile  :  in  the 
vegetable  kingdom,  a  majority  may  have  a  certain  but  very  low 
degree  of  fertility ;  but  this  is  also  the  case  in  many  unions 


May,  1869,  5)  as  the  common  designation  of  any  group  or  collection  of  indi- 
viduals whose  characters  are  continued  through  successive  generations, 
whether  it  be  permanent  variety,  subspecies,  species,  or  group  consisting 
of  very  similar  species,  the  term  not  implying  any  decision  of  this  question. 
If  this  use  of  the  term  race  prevails,  Subspecies  will  probably  take  its  place 
as  the  designation  of  the  highest  grade  of  variety.  The  objection  to  this  is 
that  the  subspecific  and  specific  names  would  be  more  liable  to  be  confused. 
1  Half-breed  is  a  common  equivalent  term  in  the  animal  kingdom :  Latin, 
Mistua  or  Mixtus  ;  French,  Metis. 

II 


322  TAXONOMY. 

within  the  species,  and  especially  in  the  application  of  the  pollen 
to  the  stigma  of  the  same  blossom.  Commonly  the  sterility  of 
hybrids  is  owing  to  the  impotence  of  the  stamens,  which  perfect 
no  pollen  ;  and  most  such  hybrids  may  be  fertilized  by  the  pollen 
of  the  one  or  the  other  parent.  Then  the  offspring  either  in 
the  first  or  second  generation  reverts  to  the  fertilizing  species. 
Moreover,  certain  hybrids,  such  as  those  of  Datura,  which  are 
fully  fertile  per  se,  divide  in  the  offspring,  partly  in  the  first  gen- 
eration, and  completely  in  two  or  three  succeeding  generations, 
into  the  two  component  species,  even  when  close- fertilized.1  (In 
part  this  may  come  from  adventive  embryo-formation,  533.) 

639.  There  appears,  therefore,  to  be  a  real  ground  in  nature 
for  species,  notwithstanding  the  difficulty  and  even  impossibility 
in  many  cases  of  defining  and  limiting  them. 

640.  Species  is  taken  as  the  unit  in  zoological  and  botanical 
classification.      Important  as  varieties   are   in  some   respects, 
especially  under   domestication  and  cultivation,  they  figure  in 
scientific   arrangement  only  as   fractions   of  species.     Species 
are  the  true  subjects  of  classification.     The  aim  of  systematic 
natural  history  is  to  express  their  relationship  to  each  other. 

641.  The  whole  ground  in  nature  for  the  classification  of  spe- 
cies is  the  obvious  fact  that  species  resemble  or  differ  from  each 
other  unequally  and  in  extremely  various  degrees.     If  this  were 
not  so,  if  related  species  differed  one  from  another  by  a  constant 
quantity,  so  that,  when  arranged  according  to  their  resemblances, 
the  first  differed  from  the  second  about  as  much  as  the  second 
from  the  third,  and  the  third  from  the  fourth,  and  so  on,  —  or  if 
the  species  blended  as  do  the  colors  of  the  rainbow,  —  then,  with 
all  the  diversit}1-  in  the  vegetable  kingdom  there  actually  is,  there 
could  be  no  natural  foundation  for  their  classification.    The  mul- 
titude of  species  would  render  it  necessary  to  classify  them,  but 
the  classification  would  be  wholly  artificial  and  arbitrary.     The 
actual  constitution  of  the  vegetable  kingdom,  however,  as  ap- 
pears from  observation,  is  that  some  species  resemble  each  other 
very  closely  indeed,  others  differ  as  widely  as  possible,  and  be- 
tween these  the  most  numerous  and  the  most  various  grades  of 


1  According  to  Naudin  in  Comptes  Kendus,  xlix.  1859,  &  Iv.  1862.  See 
also  Naudin's  memoir  on  hybridity  in  plants  in  Ann.  Sci.  Nat.  ser.  4,  xix. 
1863,  pp.  180-203,  &  in  Mem.  Acad.  Sci.  .  .  .  For  the  literature  on  vegetable 
hybrids,  see  Kcelreuter,  Nachricht,  &c.,  1761,  and  Appendices,  1763-1766; 
Herbert,  on  Amaryllidaceae,  1837  ;  C.  F.  Gzertner,  Versuche  und  Beobachtun- 
gen  ueber  die  Bastarder/eugung  in  Pflanzenreich,  1849;  Wichura,  Die 
Bastardbefruchtung  im  Pflanzenreich,  erlautertert  an  den  Bastarden  der 
W*id«n ;  and  the  memoir  of  Naudin  referred  to. 


PRINCIPLES   OF   CLASSIFICATION.  323 

resemblance  or  difference  are  presented,  but  always  with  a  mani- 
fest tendency  to  compose  groups  or  associations  of  resembling 
species, — groups  the  more  numerous  and  apparently  the  less 
definite  in  proportion  to  the  number  and  the  nearness  of  the- 
points  of  resemblance.  These  various  associations  the  naturalist 
endeavors  to  express,  as  far  as  is  necessary  or  practicable,  by  a 
series  of  generalizations,  the  lower  or  particular  included  in  the 
higher  or  more  comprehensive.  All  kinds  of  differences  are 
taken  into  account,  but  only  the  most  constant  and  definite  ones 
are  relied  on  for  characters,  i.  e.  distinguishing  marks.  Linnaeus 
and  the  naturalists  of  his  day  used  names  for  only  three  grades 
of  association,  or  groups  superior  to  species,  viz.  the  Genus,  the 
Order,  and  the  Glass  ;  and  these  are  still  the  principal  members 
of  classification. 

642.  Genera  (plural  of  Genus)  are  the  more  particular  or 
special  groups  of  related  species.  They  are  groups  of  species 
which  are  much  alike  in  all  or  most  respects,  —  which  are  con- 
structed, so  to  say,  upon  the  same  particular  model,  with  only 
circumstantial  differences  in  the  details.  They  are  not  neces- 
sarily nor  generally  the  lowest  definable  groups  of  species,  but 
*re  the  lowest  most  clearly  definable  groups  which  the  botanist 
recognizes  and  accounts  worthy  to  bear  the  generic  name ;  for 
the  name  of  the  genus  with  that  of  the  species  added  to  it  is  the 
scientific  appellation  of  the  plant  or  animal.  Constituted  as  the 
vegetable  and  animal  kingdoms  are,  the  recognition  of  genera,  or 
groups  of  kindred  species,  is  as  natural  an  operation  of  the  mind 
as  is  the  conception  of  species  from  the  association  of  like  indi- 
viduals. This  is  because  many  genera  are  so  strongly  marked, 
at  least  so  far  as  ordinary  observation  extends.  Every  one 
knows  the  Rose  genus,  composed  of  the  various  species  of  Roses 
and  Sweetbriers ;  the  Bramble  genus,  comprising  Raspberries, 
Blackberries,  &c.,  is  popularly  distinguished  to  a  certain  extent ; 
the  Oak  genus  is  distinguished  from  the  Chestnut  and  the  Beech 
genus  ;  each  is  a  group  of  species  whose  mutual  resemblance  is 
greater  than  that  of  any  one  of  them  to  any  other  plants.  The 
number  of  species  in  such  a  group  is  immaterial,  and  in  fact  is 
very  diverse.  A  genus  may  be  represented  by  a  single  known 
species,  when  its  peculiarities  are  equivalent  in  degree  to  those 
which  characterize  other  genera.  This  case  often  occurs ;  al- 
though, if  this  were  universally  so,  genus  and  species  would  be 
equivalent  terms.  If  only  one  species  of  Oak  were  known,  the 
Oak  genus  would  have  been  as  explicitly  discerned  as  it  is  now 
that  the  species  amount  to  three  hundred  ;  and  better  defined, 
for  now  there  are  forms  quite  intermediate  between  Oak  and 


324  TAXONOMY. 

Chestnut.  Familiar  illustrations  of  genera  in  the  animal  king- 
dom are  furnished  by  the  Cat  kind,  to  which  belong  the  domestic 
Cat,  the  Catamount,  the  Panther,  the  Lion,  the  Tiger,  the  Leop- 
ard, &c.  ;  and  by  the  Dog  kind,  which  includes  with  the  Dog 
the  different  species  of  Foxes  and  Wolves,  the  Jackal,  &c.  The 
languages  of  the  most  barbarous  as  well  as  of  civilized  people 
everywhere  show  that  they  have  recognized  such  groups.  Natu- 
ralists merely  give  to  them  a  greater  degree  of  precision,  and 
indicate  what  the  points  of  agreement  are. 

643.  If  most  genera  were  as  conspicuously  marked  as  those 
from  which  these  illustrations  are  taken,  genus  would  be  as  defi- 
nitely grounded  in  nature  as  species.  But  popularly  recognized 
genera,  rightly  based,  are  comparatively  few.  Popular  nomen- 
clature, embodying  the  common  ideas  of  people,  merely  shows 
that  generic  groups  are  recognizable  in  a  considerable  number 
of  cases,  but  not  that  the  whole  vegetable  or  the  whole  animal 
kingdom  is  divisible  into  a  definite  number  of  such  groups  of 
equally  or  somewhat  equally  related  species.  The  naturalist 
discerns  the  ground  of  genera  in  characters  which  the  casual  and 
ordinary  observer  overlooks  ;  and,  taking  the  idea  of  genera 
from  the  numerous  well-marked  instances  as  the  norm,  applies 
it  as  well  as  possible  to  the  less  obvious  or  less  natural  cases,  and 
groups  all  known  species  under  genera.  Resemblances  among 
the  species  when  rightly  grouped  into  genera,  though  real,  are 
often  so  unequal  in  degree,  that  certain  species  may  be  about  as 
nearly  related  to  neighboring  genera.  So  that  the  recognition  of 
genera  even  more  than  of  species  is  a  matter  of  judgment,  and 
even  of  conventional  agreement  as  to  how  and  where  a  certain 
genus  shall  be  limited,  and  what  particular  association  of  species 
shall  hold  the  position  of  genus.  All  the  species  of  a  genus  must 
accord  in  every  important  structure ;  but  extended  observation 
only  can  settle  the  question  as  to  what  are  important  and  what 
are  incidental  characters.  For  example,  the  pinnatifid  or  sinu- 
ate leaf  might  have  been  thought  as  essential  to  the  Oak  genus 
as  the  acorn-cup ;  but  many  Oaks  are  now  known  with  entire 
leaves,  resembling  those  of  Willow  or  Laurel.  An  open  acorn- 
cup  beset  with  imbricated  scales  is  a  character  common  to  all 
European  and  American  Oaks  ;  but  in  numerous  Asiatic  species 
the  cup  bears  concentric  or  spiral  lamellae  instead,  and  in  others 
the  cup  takes  the  form  of  a  naked  and  closed  sac.  Maples  have 
palmately- veined  and  lobed  leaves ;  but  one  species  has  undi- 
vided and  pinnately-veined  leaves.  The  Apple  and  the  Pear 
under  one  view  are  of  the  same  genus,  under  another  they  rep- 
resent different  genera. 


THE  PRINCIPLES  OF  CLASSIFICATION.  325 

644.  The  genus  must  be  based  on  close  relationship  of  species, 
but  not  necessarily  on   the   closest.      Raspberries  differ  from 
Blackberries,  but  must  be  ranked  in  the  same  genus  ;  and  so  of 
Plums  and  Cherries.     For  the  groups  which  are  to  bear  the 
generic  name  must  be  as  distinct  and  definite  as  possible. 

645.  Orders  are  to  genera  what  genera  are  to  species.     They 
are  groups  of  a  higher  rank  and  wider  comprehension,  expressive 
of  more  general  resemblances,  or,  in  other  language,  of  remoter 
relationship.     As  all  species  must  be  ranked  in  genera,  so  all 
genera  must  be  ranked  in  orders.     FAMILY  in  botany  is  synony- 
mous with  order :  at  least  natural  orders  and  families  (however 
distinguished  in  zoology)  have  always  in  botany  been  inter- 
changeable terms,  and  will  probably  so  continue.1 

646.  As  examples  of  orders  in  the  vegetable  kingdom  take 
the  Oak  family,  composed  of  Oaks,  Chestnuts,  Beeches,  &c. ; 
the  Pine  family,  of  Pines,  Spruces,  Larches,  Cedars,  Araucaria, 
Cypresses,  and  their  allies  ;  the  Rose  family,  in  which  Brambles, 
Strawberries,  Plums  and  Cherries,  Apples  and  Pears  are  asso- 
ciated with  the  Rose  in  one  somewhat  multifarious  order. 

647.  Classes  are  to  orders  what  these  are  to  genera.     They 
express  still  more  comprehensive  relations  of  species  ;  each  class 
embracing  all  those  species  which  are  framed  upon  the  same 
broad  plan  of  structure,  however  differently  that  plan  may  be 
carried  out  in  particulars. 

648.  Kingdom  must  be  added,  to  represent  the  highest  gener- 
alization.   All  subjects  of  biological  classification  belong  either 
to  the  vegetable  or  animal  kingdom.     Mineralogy,  Chemistry, 
&c.,  may  use  the  same  terms  (genus,  species,  &c.)  in  an  analo- 
gous way;   but  the  classification  of  substances  rests  on  other 
foundations  than  that  of  beings. 

649.  The  sequence  of  groups,  rising  from  particular  to  univer- 
sal, is  Species,  Genus,  Order,  Glass,  Kingdom;  or,  in  descending 
from  the  universal  to  .the  particular, 

KINGDOM, 
CLASS, 

ORDER, 
GENUS, 

SPECIES. 

1  Order  is  the  older  term,  and  that  which  associates  best  with  the  technical 
Latin  names.  Family  is  a  happy  term,  which  associates  itself  well  with 
English  names.  But  its  use  is  attended  with  this  incongruity,  that  the  tribe 
(653)  in  natural  history  classification  is  subordinate  to  the  family.  In 
zoology,  order  is  distinguished  from  family  as  the  next  higher  grade. 


326  TAXONOMY. 

650.  This  is  the  common  framework  of  natural  history  classi- 
fication.    All  plants  and  all  animals  belong  to  some  species ; 
every  species  to  some  genus ;    every  genus  to  some  order  or 
family ;  every  order  to  some  class ;  every  class  to  one  or  the 
other  kingdom.1    But  this  framework,  although  all  that  is  re- 
quisite in  some  parts  of  natural  history,  does  not  express  all  the 
observable  gradations  of  relationship  among  species.     And  even 
gradations  below  species  have  sometimes  to  be  classified.     The 
series  is  capable  of  extension ;  and  extension  is  often  requisite 
on  account  of  the  large  number  of  objects  to  be  arranged,  and 
the  various  degrees  of  relationship  which  may  come  into  view. 

651.  This  is   effected  by  the   intercalation   of  intermediate 
grades,  to  be  introduced  into  the  system  only  when   there  is 
occasion  for  them.     And  in  botany  one  or  more  grades  superior 
to  the  classes  are  needful;  for  first  and  foremost  is  the  great 
division  of  all  plants  into  a  higher  and  a  lower  SERIES  2  (or  sub- 
kingdom)  ,  the  Phaenogarnous  and  the  Cryptogamous. 

652.  The  grades  intercalated  into  the  long-established  sequence 
of  Class,  Order,  Genus,  and  Species,  with  new  names,  are  mainly 
two,  Tribe  and  Cohort. 

653.  Tribe  has  been  for  a  generation  or  two  thoroughly  estab- 
lished in  both  kingdoms,  as  a  grade  inferior  to  order  and  supe- 
rior to  genus.     In  botanical  classification,  much  use  is  made  of 
this  grade,  genera  being  grouped  into  tribes. 

654.  Cohort  (Lat.  Gohors)  is  of  more  recent  introduction,  at 
least  in  Botany,  but  is  becoming  established  for  a  grade  next 
above  that  of  order.    Orders  are  grouped  into  cohorts.    Lindley 
hit  upon  a  good  English  name  for  this  grade,  that  of  Alliance. 
But  this  word  has  no  available  Latin  equivalent ;  while  cohort 
takes  equally  well  a  Latin  or  an  English  form. 

655.  Finally,  each  grade  is  capable  of  being  doubled  by  the 
recognition  of  one  like  it  and  immediately  subordinate  to  it,  and 
with  designation  directly  expressive  of  the  subordination.     For 

1  Not  recognizing  HaeckeFs  third  kingdom  of  Protista,  consisting  of  those 
lowest  forms  of  being  from  which  the  animal   and   vegetable  kingdoms 
emerge. 

2  Answering  to  the  French  Embranchement  in  zoology.    For  this  it  is  pro- 
posed to  use  the  word  Division  (Divisio)  :  see  Laws  of  Botanical  Nomencla- 
ture adopted  by  the   International  Botanical  Congress  held   at  Paris  in 
August,  1867  ;  together  with  a  Historical  Introduction  and  a  Commentary,  by 
Alph.  DeCandolle,  —  English  translation,  London,  1868  ;  the  original  French 
edition,  Paris,  1867.     Perhaps  no  better  name  can  be  found ;  but  the  elder 
DeCandolle  brought  Divisio  into   common  use  for  a  grade  subordinate  to 
tribe.    Endlicher  employed  the  term  Regio.    We  have  used  Series,  and 
much  prefer  it 


THE  PRINCIPLES   OF   CLASSIFICATION.  327 

example,  if  Dicotyledones  and  Monocotyledones  be  the  two 
classes  of  Phaenogamia,  the  former  (and  only  the  former)  is 
divided  upon  very  important  characters  into  two  branches,  of  far 
higher  rank  than  the  cohorts,  viz.  the  Angiospermae  and  the 
Gymnospermse,  which  take  the  name  of  SUBCLASSES.  Orders, 
especially  the  more  comprehensive  ones,  often  comprise  two  or 
more  groups  so  distinct  that  it  may  fairly  be  a  question  whether 
they  are  not  of  ordinal  rank  :  such  take  the  name  of  SUBORDERS. 
Tribes  in  like  manner  may  comprise  groups  of  similar  relative 
value  :  these  are  SUBTRIBES.  Genera  may  comprise  sections  of 
species  which  might  almost  as  well  rank  as  genera  themselves  : 
to  mark  their  importance  and  pretension  (which  may  come  to 
be  allowed) ,  they  are  termed  SUBGENERA.  Finally,  forms  which 
are  ranked  as  varieties,  but  which  may  establish  a  claim  to  be 
distinct  species,  are  sometimes  termed  SUBSPECIES.  Even  what 
we  regard  as  a  variety  may  comprise  more  or  less  divergent 
forms,  to  be  distinguished  as  SUB  VARIETIES. 

656.  Some  of  the  larger  and  most  diversified  orders,  tribes, 
genera,  or  species  may  require  all  these  analytical  appliances, 
and  even  more,  for  their  complete  elucidation ;    while  others, 
comparatively  homogeneous,  offer  no  ground  for  them.      But 
when  these  grades,  or  some  of  them,  come  into  use,  they  are 
always  in  the  following  sequence :  — 

KINGDOM, 

SERIES  or  Division,  or  Sub-kingdom, 
CLASS, 

Subclass, 
Cohort, 

ORDER  or  Family, 
Suborder, 
TRIBE, 
Subtribe, 
GENUS, 

Subgenus, 
Section, 

Subsection, 
SPECIES, 

Subspecies  or  Race, 
Variety, 

Subvariety. 

657.  Nature  and  Meaning  of  Affinity.    These  grades,  the  higher 
including  the  lower,  denote  degrees  of  likeness   or  difference. 
Plants  belonging  respectively  to  the  two  great  series  or  primary 
divisions  may  accord  only  in  the  most  general  respects,  in  that 
which  makes  them  plants  rather  than  animals.     Plants  of  the 
same  variety  are  generally  as  much  alike  as  if  they  were  of  the 


328  TAXONOMY. 

same  immediate  parentage.  All  plants  of  the  same  species  are 
so  much  alike  that  they  are  inferred  to  have  descended  from 
a  common  stock,  and  their  differences,  however  grave,  are  sup- 
posed to  have  arisen  from  subsequent  variation,  and  the  more 
marked  differences  to  have  become  fixed  through  heredity.  This 
is  included  in  the  idea  of  species.  Descent  from  a  common 
origin  explains  the  likeness,  and  is  the  only  explanation  of  it. 

658.  But  what  is  the  explanation  of  the  likeness  between  the 
species  themselves?  As  respects  nearly  related  species,  the  answer 
is  clear.     Except  for   practical   purposes  and  in  an  arbitrary 
way,  no  certain  and  unfailing  distinction  can  be  drawn  between 
varieties  of  the  highest  grade  and  species  of  closest  resemblance. 
It  cannot  reasonably  be  doubted  that  they  are  of  similar  origi- 
nation.    Then  there  are  all  gradations  between  very  closely  and 
less  closety  related  species  of  the  same  genus  of  plants. 

659.  The  Theory  of  Descent,  that  is,  of  the  diversification  of  the 
species  of  a  genus  through  variation  in  the  lapse  of  time,  affords 
the  only  natural  explanation  of  their  likeness  which  has  yet  been 
conceived.     The   alternative  supposition,  that  all  the  existing 
species  and  forms  were  originally  created  as  they  are,  and  have 
come  down  essentially  unchanged  from  the  beginning,  offers  no 
explanation  of  the  likeness,  and  even  assumes  that  there  is  no 
scientific  explanation  of  it.     The  hypothesis  that  the  species  of 
a  genus  have  become  what  they  are  by  diversification  through 
variation  is  a  very  old  one  in  botany,  and  has  from  time  to  time 
been  put  forward.     But  until  recently  it  has  had  little  influence 
upon  the  science,  because  no  clear  idea  had  been  formed  of  any 
natural  process  which  might  lead  to  such  result.     Doubtless,  if 
variation,  such  as  botanists  have  to  recognize  within  the  species, 
be  assumed  as  equally  or  even  more  operative  through  long  ante- 
rior periods,  this  would  account  for  the  diversification  of  an 
original  species  of  a  genus  into  several  or  many  forms  as  differ- 
ent as  those  which  we  recognize  as  species.     But  this  would  not 
account  for  the  limitation  of  species,  which  is  the  usual  (but 
not  universal)   characteristic,  and  is  an  essential  part  of  the 
idea  of  species.     Just  this  is  accounted  for  by 

660.  Natural  Selection.     This  now  familiar  term,  proposed  by 
Charles  Darwin,  was  suggested  by  the  operations  of  breeders  in 
the  development  and  fixation  of  races  for  man's  use  or  fancy  ;  — 
in  animals  by  breeding  from  selected  parents,  and  selecting  for 
breeding  in  each  generation  those  individuals  only  in  which  the 
desired  points  are  apparent  and  predominant ;  in  the  seed-bed 
by  rigidly  destroying  all  plants  which  do  not  show  some  desirable 
variation,  breeding  in  and  in  from  these,  with  strict  selection  of 


THE  PRINCIPLES   OF   CLASSIFICATION.  329 

the  most  variant  form  in  the  particular  line  or  lines,  until  it  be- 
comes fixed  by  heredity  and  as  different  from  the  primal  stock 
as  the  conditions  of  the  case  allow.  In  nature,  the  analogous 
selection,  through  innumerable  generations,  of  the  exceedingly 
small  percentage  of  individuals  (as  ova  or  seeds)  which  ordi- 
narily are  to  survive  and  propagate,  is  made  by  competition  for 
food  or  room,  the  attacks  of  animals,  the  vicissitudes  of  climate, 
and  in  fine  by  all  the  manifold  conditions  to  which  they  are 
exposed.  In  the  Struggle  for  Life  to  which  they  are  thus  inevi- 
tably exposed,  only  the  individuals  best  adapted  to  the  circum- 
stances can  survive  to  maturity  and  propagate  their  like.  This 
Survival  of  the  Fittest,  metaphorically  expressed  by  the  phrase 
natural  selection,  is  in  fact  the  destruction  of  all  weaker  com- 
petitors, or  of  all  which,  however  they  might  be  favored  by  other 
conditions,  are  not  the  most  favored  under  the  actual  circum- 
stances. But  seedlings  varying,  some  in  one  direction,  some  in 
another,  are  thereby  adapted  to  different  conditions,  some  to  one 
kind  of  soil  or  exposure,  some  to  another,  thus  lessening  the  com- 
petition between  the  two  most  divergent  forms,  and  favoring  their 
preservation  and  farther  separation,  while  the  intermediate  forms 
perish.  Thus  an  ancestral  type  would  become  diversified  into 
races  and  species.  Earlier  variation  under  terrestrial  changes 
and  vicissitudes,  prolonged  and  various  in  geological  times  since 
the  appearance  of  the  main  types  of  vegetation,  and  the  attendant 
extinctions,  are  held  to  account  for  genera,  tribes,  orders,  &c., 
and  to  explain  their  actual  affinities.  Affinity  under  this  view 
is  consanguinity ;  and  classification,  so  far  as  it  is  natural,  ex- 
presses real  relationship.  Classes,  Orders,  Tribes,  &c.,  are  the 
earlier  or  main  and  successful  branches  of  the  genealogical  tree, 
genera  are  later  branches,  species  the  latest  definitely  developed 
ramifications,  varieties  the  developing  buds.1 

661.  Except  as  to  those  changes  in  size,  luxuriance,  or  depau- 
peration and  the  like,  in  which  plants,  especially  seedlings, 
respond  promptly  to  external  influences,  as  to  heat  or  cold, 

1  For  the  inception  of  this  theory  of  descent  in  the  form  which  has  within 
the  last  twenty  years  profoundly  affected  natural  history,  and  developed  a 
copious  literature,  see  a  short  paper  On  the  Variation  of  Organic  Beings  in 
a  State  of  Nature ;  On  the  Natural  Means  of  Selection ;  and  On  the  Com- 
parison of  Domestic  Races  and  True  Species,  by  Charles  Darwin,  also  On 
the  Tendency  of  Varieties  to  depart  indefinitely  from  the  Original  Type,  by 
Alfred  Russell  Wallace,  both  read  to  the  Linnean  Society,  July  1,  1858,  and 
published  in  its  Journal  of  the  Proceedings,  iii.  (Zoology)  45-62.  For  the 
development  of  the  doctrine,  see  Darwin's  "  Origin  of  Species  by  Means  of 
Natural  Selection,"  "  The  Variation  of  Animals  and  Plants  under  Domes- 
tication," and  various  other  works;  Wallace's  "Geographical  Distribution 
of  Animals,"  &c.  For  some  expositions,  see  Gray's  "  Darwiniana." 


330  TAXONOMY. 

moisture  or  dryness  (which  are  transient  and  comparatively 
unimportant) ,  variation,  or  the  unlikeness  of  progeny  to  parent, 
is  occult  and  inexplicable.  If  sometimes  called  out  by  the 
external  conditions,  it  is  by  way  of  internal  response  to  them. 
In  Darwin's  conception,  variation  of  itself  does  not  tend  in  any 
one  particular  direction :  he  appears  to  attribute  all  adaptation 
to  the  sorting  which  results  from  the  struggle  for  existence  and 
the  survival  of  the  fittest.  We  have  supposed,  and  Naegeli  takes 
a  similar  view,1  that  each  plant  has  an  internal  tendency  or  pre- 
disposition to  vary  in  some  directions  rather  than  others  ;  from 
which,  under  natural  selection,  the  actual  differentiations  and 
adaptations  have  proceeded.  Under  this  assumption,  and  taken 
as  a  working  hypothesis,  the  doctrine  of  the  derivation  of  species 
serves  well  for  the  co-ordination  of  all  the  facts  in  botany,  and 
affords  a  probable  and  reasonable  answer  to  a  long  series  of 
questions  which  without  it  are  totally  unanswerable.  It  is  sup- 
ported  by  vegetable  palaeontology,  which  assures  us  that  the 
plants  of  the  later  geological  periods  are  the  ancestors  of  the 
actual  flora  of  the  world.  In  accordance  with  it  we  may  explain, 
in  a  good  degree,  the  present  distribution  of  species  and  other 
groups  over  the  world.  It  rationally  connects  the  order  of  the 
appearance  of  vegetable  types  in  time  with  the  grades  of  differ- 
entiation and  complexity,  both  proceeding  from  the  simpler,  or 
lower  and  more  general,  to  the  higher  and  more  differentiated 
or  special ;  it  explains  by  inheritance  the  existence  of  function- 
less  parts  ;  throws  light  upon  the  anomalies  of  parasitic  plants  in 
their  various  gradations,  upon  the  assumption  of  the  most  various 
functions  by  morphologically  identical  organs,  and  indeed  illumi- 
nates the  whole  field  of  morphology  with  which  this  volume  has 
been  occupied.  It  follows  that  species  are  not  "  simple  curiosities 
of  nature,"  to  be  catalogued  and  described  merely,  but  that  they 
have  a  history,  the  records  of  which  are  impressed  upon  their 
structure  as  well  as  traceable  in  their  geographical  and  palaeon- 
tological  distribution.  This  view,  moreover,  explains  the  re- 
markable fact  that  the  characters  in  which  the  affinities  of 
plants  are  mainly  discerned  (and  which  therefore  serve  best 
for  orders,  tribes,  and  other  principal  groups)  are  commonly  such 
as  are  evidently  of  small  if  any  importance  to  the  plant's  well- 
being,  and  that  they  run  like  threads  through  a  series  of  species 
of  the  greatest  diversity  in  habit,  mode  of  life,  and  particular 
adaptations  to  conditions.2 

1  Entstehung  und  Begriff  der  naturhistorischen  Art.   Zweite  Auflage,  1865. 

2  This  is  a  corollary  of  natural  selection,  which  can  take  effect  only 
upon  useful  characters,  i.  e.  upon  structures  which  play  some  active  part 


BOTANICAL   CLASSIFICATION.  331 

662.  The  fixity  of  species  under  this  view  is  not  absolute  and 
universal,  but  relative.     Not,  however,  that  specific  changes  are 
necessitated  in  virtue  of  any  fixed  or  all-controlling  natural  law. 
Some  of  the  lowest  forms  have  existed  essentially  unchanged 
through  immense  geological  periods  down  to  the  present  time ; 
some  species  even  of  trees  are  apparently  unchanged  in  the  lapse 
of  time  and  change  of  conditions  between  the  later  tertiary  period 
and  our  own  day,  during  which  most  others  have  undergone 
specific  modification.     Such  modifications  are  too  slow  to  effect 
in  any  wise  the  stability  and  practical  application  of  botanical 
classification. 

SECTION  II.     BOTANICAL  CLASSIFICATION. 

663.  Natural  and  Artificial  Classifications  may  be  distinguished. 
A  natural  classification  in  botany  aims  to  arrange  all  known  plants 
into  groups  in  a  series  of  grades  according  to  their  resemblances, 
and  their  degrees  of  resemblance,  in  all  respects,  so  that  each 
species,  genus,  tribe,  order,  &c.,  shall  stand  next  to  those  which 
it  most  resembles  in  all  respects,  or  rather  in  the  whole  plan  of 
structure.     For  two  plants  may  be  very  much  alike  in  external 
appearance,  yet  very  different  in  their  principal  structure.    Arti- 
ficial classifications  single  out  one  or  more  points  of  resemblance 
or  difference  and  arrange  by  those,  without  reference  to  other 
considerations,  convenience   and  facility  being  the   controlling 
principles.     The  alphabetical  arrangement  of  words  in  a  dic- 
tionary, and  the  sexual  system  in  botany  by  Linnaeus  (or  rather 
a  part  of  it) ,  —  in  which  plants  are  arranged  in  classes  upon  the 
number  of  their  stamens,  and  in  orders  upon  the  number  of 
pistils,  — are  examples  of  artificial  classification.     The  arrange- 
ment of  the  words  of  a  language  under  their  roots,  and  with  the 
derivative  under  the  more  primitive  forms,  would  answer  to  a 
natural  classification. 

in  the  life  of  the  plant,  and  which  therefore  undergo  modification  under 
changing  conditions.  Unessential  structures  accordingly  are  left  unaltered 
or  are  only  incidentally  modified.  And  so  these  biologically  unessential 
points  of  structure,  persisting  through  all  adaptive  changes,  are  the  clews 
to  relationship.  Thus,  Rubiaceae  are  known  by  insignificant  stipules,  Ano- 
naceae  by  ruminated  albumen,  Rhamnaceae  by  a  valvate  calyx  and  stamens 
before  the  petals,  &c.  Paradoxical  as  it  may  seem,  it  is  not  although,  but 
because,  they  are  of  small  biological  importance  that  they  are  of  high  clas- 
sificatory  (i.  e.  of  genealogical)  value. 

On  considerations  like  these,  characters  are  divided  into  adaptive  or  bio- 
logical on  the  one  hand,  and  genealogical  or  genetic  on  the  other.  The  saga- 
cious naturalist  seizes  upon  the  latter  for  orders  and  the  like ;  while  the 
former  are  prominent  in  genera,  &c. 


332  TAXONOMY. 

664.  No  artificial  classification  of  plants  could  fail  to  be 
natural  in   some  portions  and  some  respects  ;    because  plants 
which  agree  in  any  point  of  structure  likely  to  be  used  for  the 
purpose  will  commonly  agree  in  other  and  perhaps  more  impor- 
tant characters.    On  the  other  hand,  no  natural  classification  can 
dispense  with  artificial  helps  ;  nor  can  it  express  in  lineal  order, 
or  in  any  other  way,  all  the  various  relationships  of  plants,  even 
if  these  were  fully  determined  and  rightly  subordinated.     Natu- 
ralists now  endeavor  to  make  classification  as  natural  as  possible  ; 
that  is,  to  base  it  in  every  grade  upon  real  relationships.     What 
real  relationships  are,  and  how  to  express  them  in  a  general 
system  and  throughout  its  parts,  has  been  the  task  of  the  leaders 
in  botany  from  the  beginning  of  the  science  until  now  ;  and  the 
work  is  by  no  means  completed. 

665.  Linnaeus  was  perhaps  the  first  botanist  to  distinguish 
clearly  between  a  natural  and  an  artificial   classification.     He 
labored  ineffectually  upon  a  natural  classification  of  the  genera 
of  plants  into  orders  ;  and  he  devised  an  effective  artificial  classi- 
fication, which  became  so  popular  that  it  practically  superseded 
all  others  for  more  than  half  a  century,  and  has  left  a  permanent 
impression  upon  the  science.     The  last  generation  of  botanists 
who  were  trained  under  it  has  not  quite  passed  away. 

666.  Ante-Linnsean  Classification.    Linnaeus,  in  his  Philosophia 
Botanica,  divided  systematists  into   heterodox   and   orthodox : 
the  former,  those  who  classify  plants  by  their  roots,  herbage,  time 
of  flowering,  place  of  growth,  medical  and  economical  uses,  and 
the  like  ;  the  latter,  by  the  organs  of  fructification.    It  is  remark- 
able that  all  the  orthodox  or  scientific  classifications  anterior  to 
Linnaeus  made  a  primary  division  of  the  vegetable  kingdom  into 
Trees  and  Herbs,  referring  the  larger  shrubs  to  the  former  and 
the  under-shrubs  to  the  latter,  —  an  arrangement  which  began 
with  Theophrastus  and  was  continued  by  Ray  and  Tournefort. 

667.  The  three  most  important  names  in  botanical  taxonom}r 
anterior  to  Linnaeus  are  those  o'Cesalpini,  Ray,  and  Tournefort. 
Scientific  botany  commenced  with  the  former,  in  Italy,  in  the 
latter  half  of  the  sixteenth  century.    He  first  used  the  embryo  and 
its  cotyledons  in  classification,  distinguished  differences  in  the  in- 
sertion of  floral  parts,  and,  indeed  (excepting  the  primary  division 
into  trees  and  herbs) ,  founded  all  principal  characters  upon  the 
organs   of  fructification,  especially   upon   the   fruit   and  seed. 
Conrad  Gesner  of  Zurich  had  somewhat  earlier  recognized  this 
principle,  but  Cesalpini  first  applied  it. 

668.  A  century  later  (1690-99)  this  principle  was  carried  into 
practice  by  Rivinus  (a  name  latinized  from  Bachmann),  of 


BOTANICAL  CLASSIFICATION.  333 

Leipsic,  in  a  wholly  artificial  classification  founded  on  the  corolla. 
His  contemporary  in  England,  Robert  Morison,  somewhat  earlier 
began  the  publication  of  his  great  work,  the  Universal  History 
of  Plants.  In  this  was  first  attempted  a  grouping  of  plants  into 
what  are  now  called  natural  orders  ;  and  these  were  defined,  some- 
what loosely,  some  by  their  fruit,  inflorescence,  and  flowers,  others 
by  their  stems,  the  nature  of  their  juice,  &c.  But  the  two  great 
systematists  of  the  time,  who  together  laid  the  foundations  of 
modern  scientific  botany,  were  John  Ray  in  England  and  Joseph 
Pitton  de  Tournefort  in  France. 

669.  Ray's  method  of  classification  was  sketched  in  1682,  and 
was  anterior  to  Tournefort's,  but  was  amended  and  completed  in 
1703.     The  leading  fault  of  both  was  the  primary  division  into 
trees  and  herbs.     The  great  merit  of  Ray  was  his  division  of 
herbs  into  Flowerless  and  Flowering,  and  the  latter  into  Dicotyle- 
donous and  Monocotyledonous.     These  great  classes  he  divided 
and  subdivided,  by  characters  taken  from  the  organs  of  fructi- 
fication, into  what  we  should  call  natural  orders  or  families,  but 
which  he  unfortunately  called  genera.     He  noted  the  coincidence 
of  nerved  leaves  with  the  monocotyledonous  embryo,  although 
he  did  not  notice  that  his  first  division  of  arborescent  plants  was 
monocotyledonous  ;  and  he  had  a  clear  apprehension  of  genera. 

670.  Tournefort's   method  was  published   in  French  in   the 
year  1694,  in  Latin  in  1700.     It  is  more  definite  but  more  arti- 
ficial than  that  of  Ray,  being  founded  like  that  of  Rivinus  almost 
wholly  upon  modifications  of  the  corolla,  and  it  overlooked  the  dis- 
tinction between  monocotyledonous  and  dicotyledonous  embryos. 
Its  great  merit  is  that  here  genera,  as  we  now  understand  them, 
are  first  established  and  defined,  and  all  the  species  then  known 
referred  to  them ;  so  that  Tournefort  was  justly  said  by  Linnaeus 
to  be  the  founder  of  genera.    Ray  may  be  said  to  have  indicated 
the  primary  classes,  Jussieu  (in  the  next  century)  to  have  estab- 
lished natural  orders,  and  Tournefort  to  have  given  to  botany 
the  first  Genera  Plantarum. 

671.  Linnaean  Classification.     Linnaeus,  the  great  reformer  of 
botany  in  the  eighteenth  century,  thoroughly  revised  the  principles 
of  classification,  established  genera  and  species  upon  a  more  scien- 
tific basis,  and,  in  designating  species  by  a  word  instead  of  a 
descriptive   phrase,  introduced   binomial   nomenclature.  (704.) 
He  likewise  established   for  the  stamens,  and  indeed  for  the 
pistils  also,  their  supreme  importance  in  classification  (probably 
without  knowledge  of  the  clear  suggestion  to  this  effect  made 
by  Burckhard  in  a  letter  to  Leibnitz,  printed  in  1702)  ;  their 
functions,  so  long  overlooked,  being  now  ascertained.     He  also 


334  TAXONOMY. 

drew  a  clear  and  practical  distinction  between  natural  and  arti- 
ficial classifications  (663),  and  deferring  all  endeavors  to  make 
the  former  available,  except  for  genera,  he  devised  a  practical 
substitute  for  it,  as  a  key  to  the  genera,  viz.  his  celebrated 

672.  Sexual  System,  or  arrangement  of  the  genera  under  arti- 
ficial classes  and  orders,  founded  upon  the  stamens  and  pistils. 
Although  now  out  of  use,  this  artificial  classification  has  been  so 
popular  and  influential,  and  has  left  so  deep  an  impression  upon 
the  science  and  especially  upon  the  language  of  botany,  that  it 
needs  to  be  presented.     The  primary  divisions  are  the  classes, 
twenty-four  in  number.     But  the  24th  class,  Cryptogamia,  con- 
sists of  plants  which  have  not  stamens  and  pistils  and  conse- 
quently no  proper  flowers,  and  is  therefore  the  counterpart  of  the 
remaining  twenty-three  classes,  to  which  the  corresponding  name 
of  Phanerogamia  or,  in  shorter  form,  Phsenogamia  (Phsenogamous 
plants)  has  since  been  applied.     These  twenty-three  classes  are 
characterized  by  certain  modifications  and  associations  of  the 
stamens,  and  have  substantive  names,  of  Greek  derivation,  ex- 
pressive of  their  character.     The  first  eleven  comprise  all  plants 
with  perfect  (i.  e.  hermaphrodite)  flowers,  and  with  a  definite 
number  of  equal  and  unconnected  stamens.     They  are  distin- 
guished by  the  absolute  number  of  these  organs,  and  are  desig- 
nated by  names  compounded  of  Greek  numerals  and  the  word 
andria  (from  dvrJQ) ,  which  is  used  metaphorically  for  stamen,  as 
follows :  — 

Class  1.  MONANDRIA  includes  all  such  plants  with  one  stamen  to  the  flower; 

as  in  Hippuris. 

2.  DIANDRIA,  those  with  two  stamens,  as  in  the  Lilac. 
8.  TKIANDRIA,  with  three  stamens,  as  in  the  Valerian  and  Iris. 
4.  TETRANDRIA,  with  four  stamens,  as  in  the  Scabious. 
6.  PENTANDRIA,  with  five  stamens,  the  most  frequent  case. 

6.  HEXANDRIA,  with  six  stamens,  as  in  the  Lily  Family,  &c. 

7.  HEPTANDRIA,  with  seven  stamens,  as  in  Horsechestnut. 

8.  OCTANDRIA,  with  eight  stamens,  as  in  Evening  Primrose  and  Fuchsia. 

9.  ENNEANDRIA,  with  nine  stamens,  as  in  the  Rhubarb. 

10.  DECANDRIA,  with  ten  stamens,  as  in  Rhododendron  and  Ealmia. 

11.  DODECANDRIA,  with  twelve  stamens,  as  in  Asarum  and  the  Migno- 

nette ;  extended  also  to  include  those  with  from  thirteen  to  nine- 
teen stamens. 

673.  The  two  succeeding  classes  include  plants  with  perfect 
flowers  having  twenty  or  more  unconnected  stamens,  which,  in 

12.  ICOSANDRIA,  are  inserted  on  the  calyx  (perigynous),  as  hi  the  Rose 

Family ;  and  in 

la  POLYANDRIA,  on  the  receptacle  (hypogynous),  as  in  the  Buttercup, 
Anemone,  Sfc. 


BOTANICAL   CLASSIFICATION.  335 

674.  Their  essential  characters  are  not  indicated  by  their 
names  :  the  former  merely  denoting  that  the  stamens  are  twenty 
in  number ;  the  latter,  that  they  are  numerous.  —  The  two  fol- 
lowing classes  depend  upon  the  relative  length  of  the  stamens, 
namely :  — 

14.  DIDYNAMIA,  including  those  with  two  long  and  two  short  stamens, 
as  in  the  majority  of  flowers  with  bilabiate  corolla. 

16.  TETRADYNAMIA,  those  with  four  long  and  two  short  stamens,  as  in 
flowers  with  cruciferous  corolla. 

675.  These  names  signify  in  the  former  that  two  stamens,  and 
in  the  latter  that  four  stamens,  are  most  powerful.  — The  four 
succeeding  are  founded  on  the  connection  of  the  stamens,  viz. :  — 

16.  MONADELPHIA  (meaning  a  single  fraternity),  with  the  filaments 

united  in  a  single  set,  tube,  or  column,  as  in  the  Mallow. 

17.  DIADELPHIA  (two  fraternities),  with  the  filaments  united  in  two 

sets  or  parcels,  as  in  Corydalis  and  in  many  Leguminosae. 

18.  POLYADELPHIA  (many  fraternities),  with  the  filaments  united  in 

more  than  two  sets  or  parcels,  as  in  Hypericum. 

19.  SYNGENESIA   (from   Greek   words   signifying  to  grow   together), 

with  the  anthers  united  in  a  ring  or  tube,  as  in  the  Sunflower 
and  all  Compositae. 

676.  The  next  class,  as  its  name  denotes,  is  founded  on  the 
union  of  the  stamens  to  the  style  :  — 

20.  GYNANDRIA,  with  the  stamens  and  styles  consolidated,  as  in  Cypri- 

pedium  and  all  the  Orchis  Family. 

677.  In  the  three  following  classes,  the  stamens  and  pistils 
occupy  separate  blossoms  :  — 

21.  MONCECIA  (one  household)  includes  all  plants  where  the  stamens 

and  pistils  are  in  separate  flowers  on  the  same  individual ;  as  in 
the  Oak  and  Chestnut. 

22.  DICECIA  (two  households),  where  they  occupy  separate  flowers  on 

different  individuals ;  as  in  the  Willow,  Poplar,  Moonseed,  &c. 

23.  POLYGAMIA,  where  the  stamens  and  pistils  are  separate  in  some 

flowers  and  associated  in  others,  either  on  the  same  or  two  or 
three  different  plants  ;  as  in  most  Maples. 

678.  The  remaining  class  is  essentially  flowerless  ;  or  rather  its 
organs  of  reproduction  are  more  or  less  analogous  to,  but  not 
homologous  with,  stamens  and  pistils.     But,  although  Linnaeus 
suspected  a  sexuality  in  Ferns,  Mosses,  Algae,  &c.,  there  was  no 
proof  of  it  in  his  day.     So  he  named  the  class,  containing  these, 

24.  CRYPTOGAMIA,  meaning  clandestine  marriage,  the  sexes,  if  existent, 

hidden  from  view. 

679.  The  characters  of  the  classes  may  be  presented  at  one 
view,  as  in  the  subjoined  table :  — 


TAXONOMY. 


BOTANICAL  CLASSIFICATION.  337 

680.  The  orders,  in  the  first  thirteen  classes  of  the  Linnaean 
artificial  system,  depend  on  the  number  of  styles,  or  of  the 
stigmas  when  the  styles  are  wanting ;  and  are  named  by  Greek 
numerals  prefixed  to  the  word  gynia^  used  metaphorically  for 
pistil,  as  follows  :  — 

Order  1.  MONOGYNIA,  those  with  one  style  or  sessile  stigma  to  the  flower. 

2.  DIGYNIA,  those  with  two  styles  or  sessile  stigmas. 

3.  TRIGYNIA,  those  with  three  styles. 

4.  TETRAGYNIA,  those  with  four  styles. 
6.  PENTAGYNIA,  those  with  five  styles. 

6.  HEXAGYNIA,  those  with  six  styles. 

7.  HEPTAGYNIA,  those  with  seven  styles. 

8.  OCTOGYNIA,  those  with  eight  styles. 

9.  ENNEAGYNIA,  those  with  nine  styles. 

10.  DECAGYNIA,  those  with  ten  styles. 

11.  DODECAGYNIA,  those  with  eleven  or  twelve  styles. 

12.  POLYGYNIA,  those  with  more  than  twelve  styles. 

681.  The  orders  of  class  14,  Didynamia,  are  only  two  and  are 
founded  on  the  pericarp,  namely  :  — 

1.  GYMNOSPERMIA,  meaning  seeds  naked,  the  achenia-like  fruits  of  a 

4-parted  pericarp  having  been  taken  for  naked  seeds. 

2.  ANGIOSPERMIA,  with  the  seeds  evidently  hi  a  seed-vessel  or  peri- 

carp, i.  e.  the  pericarp  undivided. 

682.  The   15th   class,   Tetradynamia,   is   also    divided    into 
two  orders,  which  are  distinguished  merely  by  the  form  of  the 
pod:  — 

1.  SILICULOSA;  the  fruit  a  silicle  (561),  or  short  pod. 

2.  SILIQUOSA;  fruit  a  silique  (561),  or  more  or  less  elongated  pod. 

683.  The  orders  of  the  16th,  17th,  18th,  20th,  21st,  and  22d 
classes  depend  merely  on  the  number  of  stamens  ;  that  is,  on  the 
characters  of  the  first  thirteen  classes,  whose  names  they  likewise 
bear:   as   MONANDRIA,  with  one  stamen,  DIANDEIA,  with  two 
stamens ;  and  so  on. 

684.  The  orders  of  the  19th  class,  Syngenesia,  are  six,  namely : 

1.  POLYGAMIA  ;EQUALIS,  where  the  flowers  are  in  heads  (the  so-called 

compound  flower),  and  all  hermaphrodite. 

2.  POLYGAMIA  SUPERFLUA,  the  same  as  the  last,  except  that  the  rays, 

or  marginal  flowers  of  the  head,  are  pistillate  only. 

3.  POLYGAMIA  FRDSTRANEA,  those  with  the  marginal  flowers  neutral, 

the  others  perfect. 
4   POLYGAMIA  NECESSARIA,  where  the  marginal  flowers  are  pistillate 

and  fertile,  and  the  central  staminate  and  sterile. 
6.  POLYGAMIA  SEGREGATA,  where  each  flower  of  the  head  [or  glom- 

erule]  has  its  own  proper  involucre. 
6.  MONOGAMIA,  where  solitary  flowers  (that  is,  not  united  into  a  head) 

have  united  anthers,  as  in  Lobelia. 


338  TAXONOMY. 

685.  The  23d  class,  Polygamia,  has  three  orders,  two  of  them 
founded   on  the  characters  of  the   two  preceding  classes  and 
bearing  their  names,  and  the  third  named  upon  the  same  prin- 
ciple, namely :  — 

1.  MONCECIA,  where  both  separated  and  perfect  flowers  are  found  in 

the  same  plant. 

2.  DICECIA,  where  they  occupy  two  different  plants. 

3.  TRKECIA,  where  one  individual  bears  the  perfect,  another  the  stami- 

nate,  and  a  third  the  pistillate  flowers. 

686.  The  orders  of  the  24th  class,  Cryptogamia,  the  Flower- 
less  Plants,  are  so  man}'  natural  orders,  and  are  not  definable 
by  a  single  character.     They  are  :  — 

1.  FILICES,  the  Ferns. 

2.  Musci,  the  Mosses. 

3.  ALG-S,  which,  as  left  by  Linnaeus,  comprised  the  Hepaticae,  Lichens, 

&c.,  as  well  as  the  seaweeds. 

4.  FUNGI,  Mushrooms,  &c. 

687.  In  its  day,  this  artificial  system  well  fulfilled  its  purpose, 
and  was  preferred  to  all  others  on  the  score  of  facility  and  defi- 
niteness.     Now  no  botanist  would  think  of  employing  it,  nor 
would  it  be  chosen  for  a  key  to  genera,  which  was  its  only  legiti- 
mate use. 

688.  The  Natural  System  was  rightly  appreciated  by  Linnaeus, 
who  pronounced  it  to  be  the  first  and  last  desideratum  in  syste- 
matic botany ;  and  he  early  attempted  to  collocate  most  known 
genera  under  natural  orders  (e.  g.  Piperitce,  Palmce,  Scitamince, 
Orchidea,  Amentacece,  &c.,  sixty-seven  in  number,  including  his 
four  cryptogamic  orders) ,  but  without  definition  or  arrangement. 
In  his  later  years,  he  was  unable  to  accomplish  any  thing  more. 
The  difficult  problem  was  taken  up  by  Linnseus's  contemporary 
and  correspondent,  Bernard  de  Jussieu,  who  planted  the  botanic 
garden  at  Trianon  with  plants  grouped  into  natural  orders,  but 
published   nothing.     His  pupil,  Adanson,  who  when  a  young 
man  lived  for  several  years  in  Senegal,  and  who  was  as  remark- 
able for  eccentricity  as  for  erudition  and  ability,  published  in 
1763,  in  his  Families  des  Plantes,  the  first  complete  system  of 
natural   orders.      But  he  seems  to  have  taken  little  from   his 
teacher,  and  with  all  his  genius  to  have  contributed  little  to  the 
advancement  of  the  natural  system. 

689.  Antoine  Laurent  de  Jussieu,  nephew  of  Bernard,  has 
been  called  the  founder  of  the  natural  system  of  botany,  and  to 
him   more   than   to  any  other  one  person  this  honor   may  be 
ascribed.     In  his  Genera  Plantarum  secundum  Ordines  Natu- 

disposita,  1789,  natural  orders  of  plants,  one  hundred 


BOTANICAL  CLASSIFICATION.  339 

in  number,  were  first  established  and  defined  by  proper  char- 
acters, and  nearly  all  known  genera  arranged  under  them.  His 
primary  division  of  the  Vegetable  kingdom  was  into  Acotyledones, 
Monocotyledones,  and  Dicotyledones,  adopted  from  Ray,  with  a 
change  which  was  no  improvement.  For  his  Acotyledones,  the 
Cr3rptogamia  of  Linnaeus,  are  the  "plants  without  flowers"  of 
Ray:  they  are,  to  be  sure,  destitute  of  cotyledons  (though  not  in 
the  manner  of  Cuscuta) ,  because  destitute  of  embryo  altogether. 
The  Acotyledones  forming  his  first  class,  Jussieu  divided  the 
Monocotyledones  into  three  classes  upon  single  and  artificial 
characters,  namely  upon  the  insertion  of  the  stamens,  whether 
hypogynous,  perigynous,  or  epigynous ;  and  the  Dicotyledones, 
into  eleven  classes  on  similar  characters,  preceded  by  a  division 
into  Apetcdce,  Monopetalce,  Polypetalce,  and  Diclines  irregulares,  i.  e. 
first  upon  the  character  of  the  perianth,  then  upon  the  insertion 
of  the  stamens  or  in  Monopetalae  of  the  corolla.  The  following 
is  the  scheme  :  — 

Acotyledones CLASS  I 

(Stamens  hypogynous II. 
perigynous III. 
epigynous IV. 

(Stamens  epigynous V. 
perigynous VI. 
hypogynous VII. 

(Corolla  hypogynous VIII. 
epigynous :  anthers  separate  XI. 


(Stamens  epigynous XII. 
hypogynous  ....  XIII. 
perigynous XIV. 

Diclinous  (also  Apetalous) XV. 


690.  Auguste  Pyrame  DeCandolle  was  the  next  great  syste- 
matist.  Reversing  the  order  of  Jussieu,  who  proceeded  from  the 
lower  or  simpler  to  the  higher  or  more  complex  forms,  DeCan- 
dolle began  with  the  latter,  the  phaenogamous  or  flowering  plants, 
and  with  those  having  typically  complete  flowers.  On  account 
of  its  convenience  and  the  greater  facilities  for  studying  the  higher 
plants,  this  order  has  been  commonly  followed  ever  since.  His 
primary  division  on  anatomical  structure,  into  Vascular  and 
Cellular  plants,  was  a  backward  step,  confusing  a  portion  of  the 
lower  series  with  the  higher ;  and  the  duplicate  names  of  Exo- 
gence  and  Endogence,  appended  to  Dicotyledonese  and  Monoco- 


340  TAXONOMY. 

tyledoneae,  as  it  now  appears  should  have  been  omitted.  The 
grades  of  the  Candollean  system  superior  to  the  orders,  in  their 
final  form,  are  mainly  these  :  — 

Div.  I.  VASCULAR  (more  properly  PIL&NOGAMOUS)  PLANTS. 
CLASS  I.  DICOTYLEDONOUS  or  EXOGENOUS. 

Subclass  I.  THALAMIFLOKOCS  :  petals  (distinct)  and  stamens  on  the 

torus,  i.  e.  free. 
II.  CALTCIFLOHODS  :  petals   (distinct  or  coalescent)  and 

stamens  adnate  to  the  calyx. 

HI.  COROLLIFLOROUS  :  petals   (mostly  coalescent)  not  ad- 
nate to  calyx,  bearing  the  stamens. 
IV.  MONOCHLAMTDEOUS  :  petals  wanting. 
CLASS  II.  MONOCOTYLEDONOUS  or  ENDOGENOUS.    (No  subclasses.) 

Div.  II.  CELLULAR  (more  properly  CRYPTOGAMOUS)  PLANTS. 

CLASS  I.  JETHEOGAMOUS  :  with  sexual  apparatus,  and 
Vascular  tissue.     (Equisetacece-Filices.) 
Only  cellular  tissue.     ( Musci  and  Hepaticce. ) 

CLASS  II.  AMPHIGAMOUS  :  destitute  of  sexual  organs  and  of  other  than 
cellular  tissue.     (Lichenes,  Fungi,  Algae.) 

691.  Cryptogamous  plants  of  all  orders  are  now  known  to  be 
provided  with  sexes ;  and  the  Jussiaean  divisions  of  the  Dico- 
tyledones  into  Apetalce   (including  Diclines),  Monopetala,  and 
Polypetalce,  are  generally  preferred  to  those  of  DeCandolle.    Into 
the  present  views  of  the  classification  of  the  Cryptogamia  it  is 
unnecessary  here  to   enter.      Their  general   arrangement   into 
classes,   &c.,  is  not  yet  well  settled,  and  the  whole  taxonomy 
of  the  lower  Cryptogams  is  at  present  in  a  state  of  transition. 

692.  John  Lindley  in  successive  attempts  (between  1830  and 
1845)  variously  modified,  and  in  some  few  respects  improved, 
the  Candollean  arrangement.    But,  as  neither  his  groupings  of  the 
natural  orders  nor  the  new  classes  which  he  adopted  have  been 
approved,  his  schemes  need  not  be  here  presented.     He  must  be 
credited,  however,  with  the  first  attempt  to  carry  into  effect  a 
suggestion  made  by  Brown,  that  the  orders  should  themselves 
be  disposed  as  far  as  possible  into  superior  and  strictly  natural 
groups.     In  Llndley's  first  attempt,  such  groups  of  two  grades 
were  proposed,  the  lower  called  nixus  (tendencies),  the  higher 
cohorts.      In  his  later  and  largest  work,  The  Vegetable  King- 
dom, these  were  reduced  to  one,  and  the  name  of  alliance  was 
coined.     But  this  word  has  no  good  Latin  equivalent,  and  the 
term  cohort  (cohors)  is  preferred. 


BOTANICAL  CLASSIFICATION.  341 

693.  Robert  Brown,  next  to  Jussieu,  did  more  than  any  other 
botanist  for  the  proper  establishment  and  correct  characterization 
of  natural  orders.     Having  in  the  year  1827  published  his  dis- 
covery of  the   gymnospermy  of  Coniferae   and   Cycadaceae,  it 
was  in  Lindley's   works   that  this   was  first  turned  to  proper 
systematic  account  by  dividing  the  class  of  Dicotyledones  into 
two  subclasses,  the  Angiospermce  and  the   Gymnospermce.     The 
latter  has  been  elevated  by  the  vegetable  palaeontologists  to  the 
rank  of  a  class. 

694.  Stephen   Ladislaus  Endlicher,  of  Vienna,  a  contempo- 
rary of  Lindley,  of  less  botanical  genius,  but  of  great  erudition 
and  aptness  for  classification,  brought  out  his  complete  Genera 
Plantarum  secundum  Ordines  Naturales  disposita,  between  the 
years  1836  and  1840.     This  elaborate  work  follows  that  of  its 
predecessor,  Jussieu,  in  beginning  with  the  lower  series  of  plants 
and  ending  with  the  higher.     Its  primary  division  is  into  two 
regions :  1 .    Thallophyta,  plants  without  proper  axis  of  growth 
(developing  upward  as  stem  and  downward  as  root),  no  other 
tissue  than  parenchyma,  and  (as  was  thought)  no  proper  sexes. 
This  answers  to  the  lower  or  Amphigamous  Cellular  plants  of 
DeCandolle.  2.  Cormophyta,  plants  with  an  axis  (stem  and  root), 
with  foliage,  &c.     The  Cormophyta,  or  plants  of  the   higher 
region,  Endlicher  divided  into  three   great  sections:    1.  Acro- 
brya,  answering  to  the  higher  JEtheogamous  Cryptogamia   of 
DeCandolle,  with  which  was  wrongly  associated  a  group  of  root- 
parasitic  flowering  plants  (the  Rhizantheae)  which  were  fancied 
to  bear  spores  instead  of  embryo  in  their  seed ;  2.  Amphibrya, 
which  answer  to  Monocotyledones  ;  and,  3.  Acramphibrya,  which 
answer  to  Dicotyledons.     These  last  contain  five   cohorts:  1. 
Gymnospermece ;    2.  Apetalce ;   3.  Gamopetalce  (the  Monopetalae 
of  Jussieu  better  named)  ;   4.  Dicdypetalce  (the  Polypetalae  of 
Jussieu,  &c.).     The  cohort  in  Endlicher' s  classification,  it  will 
be  seen,  is  a  higher  grade  than  that  to  which  this  name  was 
applied  by  Lindley  in  the  more  recent  use.     For  the  latter,  i.  e. 
for  the  grade  between  these  and  the  order,  Endlicher  employee! 
the  name  of  class. 

695.  Finally,  the  Genera  Plantarum,  now  in  course  of  pub- 
lication by  George  Bentham  and  Joseph  Dalton  Hooker,  adopts 
in  a  general  way  the  Candollean  sequence  of  orders,  with  vari- 
ous emendations  ;  divides  the  class  of  Dicotyledons  into  two  sub- 
classes, Angiospermous  and  Gymnospermous ;  the  former  into 
the  Polypetalous,  Gamopetalous,  and  Apetalous  divisions ;  and 
the  first  of  these  into  the  Thalamiflorous,  Disciflorous,  and  Caly- 
ciflorous  "  series  "  (the  middle  one  composed  of  the  latter  part 


342  TAXONOMY. 

of  DeCandolle's  Thalamiflorae  with  some  of  his  Calyciflorae)  ; 
and  under  these  the  orders  are  arranged  in  cohorts,  —  fifteen 
cohorts  in  the  Polypetalae,  and  ten  under  three  "  series  "  in  the 
Gamopetalae.  The  remainder  of  this  particular  classification 
has  not  yet  appeared  in  print,  although  partly  sketched  by  its 
authors.  It  will  generally  be  adopted  in  this  country,  with  some 
occasional  minor  modifications. 

696.  Various  modifications  have  been  from  time  to  time  pro- 
posed.    One  of  the  best  of  them  in  principle  is  that  initiated  by 
Adolphe  Brongniart  and  adopted  by  many  European  botanists, 
which,  recognizing  that  most  apetalous  flowers  are  reductions  or 
degradations  of  polypetalous  types,  intercalates  the  Apetalae  or 
Monochlamydeae  among  the  Polypetalae.    But  this  has  never  yet 
been  done  in  a  satisfactory  manner,  or  without  sundering  orders 
which  should  stand  in  contiguity. 

697.  It  should  be  borne  in  mind  that  the  natural  system  of 
botany  is  natural  only  in  the  constitution  of  its  genera,  tribes, 
orders,  &c.,  and  in  its  grand  divisions  ;  that  its  cohorts  and  the 
like  are  as  yet  only  tentative  groupings ;  and  that  the  putting 
together  of  any  or  all  these  parts  in  a  system,  and  especially 
in  a  lineal  order,  necessary  as  a  lineal  arrangement  is,  must 
needs  be  largely  artificial.      So  that  even  the  best  perfected 
arrangements  must  always  fail  to  give  of  themselves  more  than 
an  imperfect  and  considerably  distorted  reflection  of  the  plan  of 
the  vegetable  kingdom,  or  even  of  our  knowledge  of  it.1 

1  In  the  first  place,  the  relationships  of  any  group  cannot  always  be 
rightly  estimated  before  all  its  members  are  known  and  their  whole  struct- 
ure understood  ;  so  that  the  views  of  botanists  are  liable  to  be  modified  with 
the  discoveries  of  every  year.  The  discovery  of  a  single  plant,  or  of  a  point 
of  structure  before  misunderstood,  has  sometimes  changed  materially  the 
position  of  a  considerable  group  in  the  system,  and  minor  alterations  are 
continually  made  by  our  increasing  knowledge.  Then  the  groups  which  we 
recognize,  and  distinguish  as  genera,  tribes,  orders,  &c.,  are  not  always,  and 
perhaps  not  generally,  completely  circumscribed  in  nature,  as  we  are  obliged 
to  assume  them  to  be  in  our  classification.  This  might  be  expected  from 
the  nature  of  the  case.  For  the  naturalist's  groups,  of  whatever  grade,  are 
not  realities,  but  ideas.  Their  consideration  involves  questions,  not  of  things, 
between  which  absolute  distinctions  might  be  drawn,  but  of  degrees  of  resem- 
blance, which  may  be  expected  to  present  infinite  gradations.  Besides,  al- 
though the  grades  of  affinity  among  species  are  most  various,  if  not  wholly 
indefinite,  the  naturalist  reduces  them  all  to  a  few,  and  treats  his  genera, 
tribes,  &c.,  as  equal  units,  or  as  distinguished  by  characters  of  about  equal 
value  throughout,  —  which  is  far  from  being  the  case.  And  in  his  works 
he  is  obliged  to  arrange  the  groups  he  recognizes  in  a  lineal  series ;  but 
each  genus  or  order,  &c.,  is  very  often  about  equally  related  to  three  or  four 
others  :  so  that  only  a  part  of  the  relationship  of  plants  can  In  any  way  be 
indicated  by  a  lineal  arrangement. 


BOTANICAL  CLASSIFICATION.  343 

698.  Even  the  great  classes  cannot  be  arranged  in  a  single 
line,  beginning  with  the  highest  Phaenogams  and  leaving  the 
lowest  in  contiguity  with  the  higher  Cryptogams.     The  Dicoty- 
ledons take  precedence  of  the  Monocotyledons  in  rank.     Yet  a 
part  of  them,  the  Gymnosperms,  are  much  the  lowest  of  all 
known  Phaenogams  as  regards   simplicity  of  floral  structure ; 
and  through  them  only  is  a  connection  with  the  higher  Cryp- 
togams to  be  traced.     The  Monocotyledons  stand  upon  an  iso- 
lated side  line,  and   have  no  such  simplified   representatives. 
In  placing  the  latter  class  between  the  Dicotyledons  and  the 
Acrogens,  the  chain  of  affinities  is  widely  sundered.     If,  yield- 
ing to  a  recent  tendency,  we  raise  the  Gymnosperms  to  the 
rank  of  a  class,  and  place  it  between  the  Monocotyledons  and 
the  Acrogens,  then  the  much  nearer  relationship  of  Gymno- 
sperms to  Angiosperms  through  Gnetaceae  and  Loranthaceae  is 
not  respected.  (606,  &c.) 

699.  Nor  can  the  angiospermous  Dicotyledons  be  disposed 
lineally  according  to  rank.      The  apetalous  and  achlamydeous 
must  be  the  lowest.     Some  are  evidently  reduced  forms  of  Poly- 
petalae  or  even  of  Gamopetalae  :  the  greater  part  cannot  without 
violence  be  thrust  into  their  ranks.    The  Gamopetalae,  especially 
those  with  much  floral  adnation,  should  represent  the  highest 
type,  the  organs  being  at  the  same  time  complete  and  most  dif- 
ferentiated from  the  foliar  state.     If  a  natural  series  could  be 
formed,  these  would  claim  the  highest  place,  with  the  Compositae 
perhaps  at  their  head.    In  the  Candollean  sequence,  they  occupy 
the  middle  ;  and  the  series  begins,  not  without  plausible  reason, 
with  orders  having  generally  complete  blossoms,  and  such  as 
most  freely  and  obviously  manifest  the  homolog}7  of  their  organs 
with  leaves,  then  rises  to  those  of  greater  and  greater  combi- 
nation and  complexit}r.  and  ends  with  those  plants  which,  with 
all  their  known  relatives,  are  most  degraded  or  simplified  by 
abortions  and  suppressions  of  parts  which  are  represented  in  the 
complete  flower.     These  are  low  in  structure,  equally  whether 
we  regard  them  as  reduced  forms  of  higher  types,  or  as  forms 
which  have  never  attained  the  full  development  and  diversifica- 
tion which  distinguish  the  nobler  orders. 

700.  Actual  classifications,  in  their  leading  features  and  in 
their  extension  to  the  cohorts,  orders,  &c.,  must  be  studied  in 
the   systematic   works  where  they  are   brought  into  use.      In 
these   are   the   applications   of  the   principles   which   are   here 
outlined.     A  separate  volume  of  this  text-book  should  illustrate 
the  structure,  relations,  and  most  important  products  of  the 
phaenogamous  natural  orders,  as  another  is  to  illustrate  the 


344  TAXONOMY. 

cryptogamous  orders.  A  synoptical  view  of  the  great  divi- 
sions only,  as  at  present  received  and  named,  is  appended. 
Definitions  and  characters  may  be  sought  in  the  present  and 
preceding  chapters. 

SERIES  L    PaffiNOGAMOUS   OR  FLOWERING  PLANTS. 
CLASS!  DICOTYLEDONS. 

SUBCLASS  L  ANGIOSPERMS. 

DlV.  1.  POLYPETALOUS. 

Div.  2.  GAMOPETALOUS.  (Monopetalous,) 
Div  3.  APETALOUS. 

SUBCLASS  IL  GYMNOSPERMS. 

ClJUW  H.  MONOCOTYLEDONS. 
Div.  1.  SPADICEOUS. 
Div.  2.  PETALOIDEOUS. 
Div.  3.  GLUMACEOUS. 

SERDZS  H.    CRYPTOGAMOUS  OR  FLOWERLESS  PLANTS. 

CLASS  m.  ACROGENS. 

Div  1.  VASCULAR.  (Ferns  and  their  allies.) 
Div.  2.  CELLULAR.  (Mosses  and  Liverworts.) 

CLASS  IV.  THALLOGENS  OR  THALLOPHYTES. 


PHYTOGRAPHY.  345 


CHAPTER    X. 

PHYTOGRAPHY. 

SECTION  I.  NOMENCLATURE. 

701.  PHYTOGRAPHY  is  the  department  of  botany  which  relates  to 
the  description  of  plants.     This  includes  names  and  terms,  also 
figures  and  signs,  as  well  as  characters  and  detailed  descriptions. 
It  comprises  two  sorts  of  names,  one  used  to  designate  organs 
or  modifications  of  organs,  the  other  to  distinguish  plants  or 
groups  of  plants.     The  former  is  Glossology  or  (to  use  the  more 
common  but  less  proper  word)  Terminology,     The  latter  is  prop- 
erly Nomenclature. 

702.  Names  of  Plants  were  at  first  only  generic  names.     The 
language  of  botan}*  being  Latin,  and  the  plants  which  the  old 
herbalists  knew  being  mostly  European,  their  scientific  names 
were  mainty  adopted  from  the  ancient  Romans  or,  through  Latin 
literature,  from  the  Greeks.     Ex.  Quercus,  Prunus,  Rosa,  Rubus, 
Trifolium ;  and  of  Latinized  Greek  names,  Agrostis,  Aristolochia, 
Colchicum,    Melilotus.  •   To    the    classical    names   others   were 
added  from  time  to  time  ;  as,  from  the  Latin,  Bidens,  ConvaUaria, 
Dentaria  ;  from  the  Greek,  Anacardium,  Glycyrrhiza,  Loranthus, 
&c.     Some  barbarous  or  outlandish  names  were  early  adopted, 
such  as  Alhagi  from  the  Arabs,  and  Adhatoda  and  Nelumbo  from 
India.     These  are  mostly  such  as  were  or  could  be  conformed  to 
Latin  ;  as  Datura  and  Ribes  from  the  Arabic,  and  later  Thcea  and 
Coff&a.     Of  American  aboriginal  names,  ffura,  Guaiacum,  and 
Yucca  are  examples.     Some  ancient  names  of  plants  commem- 
orated distinguished  men.    Ex.  Asclepias,  Euphorbia,  Lysimachia, 
Pceonia.      Tournefort    and    his    contemporaries    resumed   this 
practice,  and  named  plants  in  memory  or  in  honor  of  distin- 
guished botanists.     Ex.  Begonia,  Bignonia,  Ccesalpinia,  Fuchsia, 
Gerardia,  Lobelia,  Lonicera,  Magnolia. 

703.  When  among  plants  of  the  same  name  or  kind  different 
species  were  known,  these  were  distinguished  by  annexed   epi- 
thets.    For  example,  among  the  Pines  there  were  :  Pinus  syl- 
vestris, vulgaris  ;  Pinus  sylvestris,  montana  alter  a  ;  Pinus  sylvestris, 
montana  tertia  /  Pinus  sylvestris  maritima,  conis  firmiter  ramis 
adhcerentibus  ;  Pinus  maritima  minor  ;  Pinus  maritima  altera,  &c. 


346  PHYTOGRAPHY. 

And  as  the  number  of  known  species  increased,  so  did  the 
length  of  the  phrases  which  were  needed  for  their  discrimina- 
tion. These  "  differentiae,"  thus  used  as  specific  names  (the 
nomina  specified  of  Linnaeus) ,  became  extremely  cumbrous.  It 
was  about  in  the  middle  of  his  career  that  Linnaeus  suggested 
what  he  called  trivial  names  (nomina  trivialia)  for  the  specific 
name,  consisting  of  a  single  word ;  and  in  the  Species  Plan- 
tarum,  in  1753,  he  carried  this  idea  into  full  effect  in  Botany. 
The  step  was  a  simple  one,  but  most  important ;  and  Linnaeus 
himself,  who  generally  did  not  underrate  his  services  to  science, 
seems  hardly  to  have  appreciated  its  practical  value.1 

704.  The   Binomial   Nomenclature   in   Natural  History,  thus 
established,  first  separated  the  name  of  a  plant  or  of  an  animal 
from  its  diagnosis,  descriptive  phrase,  or  character,  and  reduced 
the  appellation  to  two  words,  the  first  that  of  its  genus,  the  sec- 
ond that  of  its  species.    The  generic  name  very  nearly  answers  to 
the  surname  of  a  person,  as  Brown  or  Jones;  the  specific  answers 
to  the  baptismal  name,  as  John  or  James.     Thus,  Quercus  alba  is 
the  botanical  appellation  of  the  "White  Oak  ;    Quercus  being  that 
of  the  genus,  and  alba  (white)  that  of  a  particular  species  ;  while 
the  Red  Oak  is  named  Quercus  rubra  ;  the  Scarlet  Oak,  Quercus 
coccinea;  the  Live  Oak,    Quercus  virens ;  the  Bur  Oak,  Quercus 
macrocarpa :   Magnolia  grandiflora  is  Large-flowered  Magnolia  ; 
M.  macrophylla,  Long-leaved  Magnolia,  and  so  on.    The  name  of 
the  genus  is  a  substantive,  or  at  least  is  a  word  taken  as  a  substan- 
tive.    That  of  a  species  is  mostly  an  adjective  adjunct,  always 
following  the  generic  name  and  in  the  same  gender.2    This  com- 
bination of  generic  and  specific  name  is  the  name  of  the  plant.8 

705.  By  this  system,  not  only  is  the  name  of  the  plant  reduced 
to  two  words,  but  a  comparatively  moderate  number  of  words 
serves  for  the  complete  designation  of  more  than  120,000  plants,4 


1  Moreover,  he  may  be  said  to  have  adopted  rather  than  originated  the 
idea ;  for  single-worded  specific  names  were  used  half  a  century  previous 
by  Bachmann,  alias  Rivinus. 

2  It  is  to  be  noted  that  the  classical  Latin  names  of  trees  are  all  feminine, 
therefore  Quercus  alba,  Pinus  rigida,  &c. 

8  The  name  of  a  subgenus  is  sometimes  written  in  between  the  two  parts 
of  the  plant's  name,  as  Prunus  (Padus)  Virginiana.  This  is  the  name  of  the 
plant  and  something  more.  In  addition  to  the  name  of  the  species,  that  of 
the  variety  or  even  subvariety  is  sometimes  added. 

*  Alphonse  DeCandolle  several  years  ago  estimated  the  known  species  of 
Flowering  Plants  at  between  100,000  and  120,000.  The  larger  number  may 
perhaps  include  the  higher  orders  of  the  Flowerless  series.  In  the  present 
state  of  our  knowledge  of  the  lower  orders  of  Cryptogams,  no  close  estimate 
can  be  well  formed  of  the  actual  number  of  species. 


NOMENCLATURE.  347 

in  a  manner  which  avoids  confusion  and  need  not  overburden  the 
memory.  The  generic  part  of  the  name  is  peculiar  to  each 
genus.  The  specific  adjunct  is  not  available  for  more  than  one 
species  of  the  same  genus,  but  may  be  used  in  any  other  genus. 
They  are  so  widely  thus  employed  that  the  number  of  specific 
may  not  exceed  that  of  generic  appellations. 

706.  To  render  this  system  of  nomenclature  most  serviceable 
for  the  ready  identification  of  such  numbers  of  plants  or  groups, 
and  for  the  clear  and  succinct  presentation  of  or  reference  to 
what  is  known  and  recorded  of  them,  rules  are  indispensable, 
and  conformity  to  admitted  rules  is  a  manifest  duty.    Such  rules 
were  systematically  formulated  first  by  Linnaeus,  in  bis  Funda- 
menta,   Critica,  and  Philosophia  Botanica,   chiefly  for  generic 
names,  some  of  them  being  of  the  nature  of  laws,  some  rather  of 
recommendations.     The  most  important  of  them  remain  in  full 
force,  while  many  of  the  more  particular  rules  restricting  the 
choice  of  names  have  been  abandoned.     The   code  was  judi- 
ciously revised   (in   his   Theorie   Elementaire)   by  DeCandolle 
"  who  was  ruled  by  the  idea  of  having  the  law  of  priority  prop- 
«rly  respected,"  was  critically  considered  by  Lindley  in  his  In- 
troduction to  Botany,   and  has  of  late  been  reformulated  by 
Alphonse  DeCandolle  under  the  sanction  of  a  Botanical  Congress 
held  at  Paris  in  1867.1 

707.  Rules  for  Naming  Plants.     These  "should  neither  be 
arbitrary  nor  imposed  by  authority.     They  must  be  founded  on 
considerations  clear  and  forcible  enough  for  every  one  to  com- 
prehend and  be  disposed  to  accept.      The  essential  point  in 
nomenclature  is  to  avoid  or  to  reject  the  use  of  forms  or  names 

1  Lois  de  la  Nomenclature  Botanique,  etc.,  Geneva  and  Paris,  1867.  In 
the  English  edition,  translated  by  Weddell :  Laws  of  Botanical  Nomenclature 
adopted  by  the  International  Botanical  Congress  held  at  Paris  in  August, 
1867,  together  with  an  Historical  Introduction  and  a  Commentary,  London, 
Reeve  &  Co.,  1868.  The  Laws,  simply,  were  reprinted  in  the  American  Journal 
of  Science  and  Arts,  July,  1868.  A  few  special  points  have  been  more  recently 
discussed  by  various  critics,  especially  in  the  Bulletin  of  the  Botanical  Society 
of  France,  and  in  that  of  the  Royal  Botanical  Society  of  Belgium.  See  like- 
wise American  Journal  of  Science  and  Arts  for  September,  1870,  and  August, 
1877  ;  also,  Bentham  in  Journal  of  the  Linnean  Society,  xvii.  189-198,  in  which 
a  just  distinction  is  indicated  between  changing  a  well-established  name  and 
giving  a  new  name  to  a  new  plant.  See  American  Journal  of  Science  for 
April,  1879. 

Mention  should  also  be  made  of  Strickland's  Report  of  a  Committee  on 
Nomenclature  to  the  British  Association  in  1842,  of  Agassiz's  classical  preface 
on  the  nomenclature  of  genera  in  his  Nomenclator  Zoologicus,  and  of  DalPs 
thorough  and  well-digested  Report  of  the  Committee  on  Zoological  Nomen. 
clature  to  the  American  Association  for  the  Advancement  of  Science,  1877, 
—  these  dealing  primarily  with  zoology. 


348  PHYTOGRAPHY. 

that  may  create  error  or  ambiguity,  or  throw  confusion  into 
science.  Next  in  importance  is  the  avoidance  of  any  useless 
introduction  of  new  names.  Other  considerations,  such  as 
absolute  grammatical  correctness,  regularity  or  euphony  of 
names,  a  more  or  less  prevailing  custom,  respect  for  persons, 
&c.,  notwithstanding  their  undeniable  importance,  are  relatively 
accessory.'  (Alph.  DeCandolle,  1.  c.) 

708.  The  following  are  universal  rules  in  scientific  nomen- 
clature :  — 

1.  Names  must  be  in  Latin  or  be  Latinized.     Those  from  the 
Greek  (which  are  more  and  more  abundant,  owing  to  the  facility 
of  this  language  for  compounding)  take  Latin  form  and  termina- 
tion.1    Those  from  modern  or  other  than  classical  languages 
should   at  least  have   a   Latin   termination.2     Hybrid  names, 
namely,  those  formed  by  the  combination  of  two  languages  (at 
least  of  Latin  and  Greek) ,  should  not  be  made.8 

2.  For  each  plant  or  group  there  can  be  only  one  valid  name, 
and  that  always  the  most  ancient,  if  it  is  tenable. 

3.  Consequently,  no  new  name  should  be  given  to  an  old  plant 
or  group,  except  for  necessity.     That  a  name  may  be  bettered 
is  no  valid  reason  for  changing  it. 

709.  Names  of  Genera  are  substantive  and  singular,  of  one 
word ;  and  the  same  name  cannot  be  used  for  two  genera  of 
plants.4    They  may  be  derived  from  any  source  whatever,  from 


1  Thus,  words  ending  with  the  Greek  os  generally  change  it  to  us,  and  with 
on  to  urn.     A  rule  not  always  observed ;  for  while  we  have  Epidendrum  and 
Oxydendrum,  Linnaeus  himself  variously  wrote  Liriodendrum  and  Liriodendron, 
Ekododendrum  and  Rhododendron ;  and  the  Greek  form  now  prevails. 

2  In  this   as  in  other  cases,  some  exceptions  are  well  established  by 
custom,  but  they  ought  not  to  be  extended.    The  rule  as  to  Latinization  is 
restricted  as  respects  orthography  by  the  necessity  of  preserving  modern 
commemorative  names  in  a  recognizable  form. 

8  But  we  cannot  change  numerous  old  names  for  this  fault,  such  as  con- 
volvuloides,  fumarioides,  ranunculoides,  and  scirpoides  (though  they  ought  to 
have  been  convolndina,  ranunculina,  and  scirpina)  ;  and  modern  botanists  have 
not  scrupled  to  append  the  expressive  and  convenient  Greek  term  -aides  (sig- 
nifying likeness)  to  generic  names  not  of  classical  origin.  Ex.  abutiloides, 
bixoides,  davallioides,  fuchsioides,  r/eniianoides,  lobelioides,  toumefortioides.  In  Eng- 
lish, some  hybrids  will  perpetuate  themselves,  as  for  instance  terminology, 
centimetre,  millimetre,  beaurocracy,  &c. 

*  Very  many,  indeed,  are  adjectives  used  as  substantives,  as  Arenaria, 
Clavaria,  Saponaria,  Tmpatiens,  Trientalis,  and  even  Gloriosa,  Mirabilis,  &c. 

Some  two-worded  generic  names  anterior  to  Linnaeus,  such  as  Dens  Leonis, 
Vitis  Idcea,  Bursa  Pastoris,  remain  for  sections  and  species,  but  not  for  gen- 
era. When  two  words  are  confluent  into  one,  they  are  not  objectionable, 
as  Laurocerasus,  Carlemania  (commemorating  Charles  Leman,  Carolus  Le- 
manus),  &c. 


NOMENCLATURE.  349 

prominent  or  peculiar  character  or  appearance,  from  localities, 
from  the  names  of  persons  (especially  of  discoverers),  from 
indigenous  or  vulgar  names,  or  even  from  arbitrary  combina- 
tions of  letters.  Unmeaning  names,  if  not  in  principle  the  best, 
are  never  misleading.  The  main  requisite  is  that  they  should 
be  euphonious,  not  too  long,  and  that  they  should  be  adaptable 
to  the  Latin  tongue.  Characteristic  names,  when  possible,  are 
among  the  best ;  such  as  Sanguinaria  for  an  herb  with  red  juice, 
Hcematoxylon  for  the  Logwood  tree,  Lithospermum  for  a  plant 
with  stony  seeds  (or  seeming  seeds) ,  Myosurus  for  a  plant  with 
gynoecium  resembling  the  tail  of  a  mouse.  Names  of  this  sort 
do  not  always  hold  out  well ;  for  Chrysanthemum,  so  called  from 
its  golden  yellow  blossoms,  now  has  many  white-flowered  species, 
Polygala  is  wholly  destitute  of  milk,  and  many  species  of  Con- 
volvulus do  not  twine.  Neat  anagrams  are  not  bad,  such  as 
Brown's  Tellima  for  a  genus  nearly  related  to  Mitella.  Personal 
generic  names  are  wholly  proper  when  dedicated  to  botanists, 
especially  to  the  discoverer  of  the  plant,  or  to  other  naturalists, 
or  to  persons  who  have  furthered  botanical  investigation  or 
exploration.  Ancient  names  of  this  kind  have  been  mentioned, 
also  some  of  those  which  commemorate  the  earlier  botanists. 
(702.)  At  present,  almost  every  devotee  of  the  science  is  thus 
commemorated,  from  Linnaeus  and  Jussieu  downward.  In 
forming  such  names,  the  name  of  the  person,  cleared  of  titles 
and  accessor}-  particles  (thus  Candollea,  not  Decandollea) ,  takes 
the  final  -a  or  -ia  and  becomes  feminine  ;  and  its  orthography  is 
preserved  as  far  as  possible,  making  only  necessary  concessions 
to  euphony  and  to  the  genius  of  the  Latin  language.1 


The  Linnaean  canon  forbade  the  use  of  the  same  generic  name  in  botany 
and  zoology,  —  a  rule  now  impossible  to  maintain.  Perhaps  we  cannot  pre- 
vent the  duplication  of  phaenogamous  names  in  the  lower  Cryptogamia. 

1  Thus,  we  may  write  Lescuria  instead  of  Lesquereuxia,  although  Michauxia. 
is  the  form  for  the  genus  dedicated  to  Michaux,  however  pronounced.  The 
genus  dedicated  to  Strangways  is  written  Stranvcesia  (although  Strangwaysia 
might  have  been  tolerable) ;  to  Andrzeiowsky,  Andreoskia ;  to  Leeuwenhoek, 
Levenhookia  (although  the  elder  DeCandolle  restored  all  the  vowels),  &c. 
As  specimens  of  overdone  simplification,  there  is  Gundelia,  named  for  Gun- 
delsheimer,  and  Goodenia,  named  for  Bishop  Goodenough,  although  Gundels- 
heimera  would  not  in  these  days  be  objected  to,  and  Goodenovia  is  faultless. 
Yet  the  names  having  been  so  introduced  into  the  science  should  remain, 
fixity  being  of  more  importance  than  perfection.  Mistaken  orthography 
of  the  name  itself  may,  however,  be  set  right.  Brown's  Lechenaultia  is  Les- 
chenaultia,  Nuttall's  Wisteria  (named  after  Dr.  Wistar)  is  Wistaria.  The 
rule  laid  down  in  the  code  as  drawn  up  by  Alphonse  DeCandolle  is; 
"  When  a  name  is  drawn  from  a  modern  language,  it  is  to  be  maintained 
just  as  it  was  made,  even  in  the  case  of  the  spelling  having  been  misunder- 


350  PHYTOGRAPHY. 

710.  The  etymology  of  a  new  genus  should  always  be  given. 
Of  the  Linnaean  restrictions,  one  holds,  viz.  that  the  names  of 
genera  are  not  to  end  in  -oides,  as  many  of  the  older  names  did. 

711.  Names  of  Species  are  commonly  and  by  preference  adjec- 
tives, agreeing  with  the  name  of  the  genus,  and  expressive  of 
some  character,  habit,  mode  or  place  of  growth,  time  of  flower- 
ing, or  commemorating  the  discoverer,  first  describer,  or  some 
one  otherwise  connected  with  its  history.     Thus,  in  the  genus 
Ranunculus,  R.  bulbosus  is  named  from  the  bulb-like  crown  or  base 
of  the  stem  ;  R .  acris,  from  the  acridity  of  the  juice ;  R.  scele- 
ratus  (the  accursed),  in  reference  to  the  same  property  ;  R .  repens, 
from  the  creeping  habit  of  the  stems  ;  R.  pusillus,  from  general 
insignificance  ;  R .  aquatilis,  from  its  growing  in  water ;  R.  ni- 
valis,  from  living  near  eternal  snow ;   R.  Pennsylvanicus,  from 
country  or  State  whence  it  was  first  made  known  to  botanists  ; 
R.  Bonplandianus,  in  honor  of  Bonpland,  one  of  the  discoverers  ; 
and  so  on.     More  commonly,  when  a  discoverer  or  investigator 
of  a  species  is  commemorated  in  the  name,  this  is  a  substantive, 
in  the  genitive,  as  Ranunculus  Nuttattii,  i.  e.  the  Ranunculus  of 
Nuttall,  instead  of  R.  Nuttallianus,  the  Nuttallian  Ranunculus. 
Yet  the  latter  form  is  preferred  when  the  species  is  named  in 
honor  of  some  one  who  did  not  discover  nor  treat  of  it  (which 
should  seldom  be)  ;  but  this  distinction  is  a  custom  rather  than 
a  rule,  and  the  form  of  the  commemorative  name  may  be  settled 
by  euphony  or  convenience.     In  any  case,  the  personal  name 
should  have  a  capital  initial. 

712.  Many  specific  names  are  substantives,  occasionally  a 
common  substantive,  as  SteUaria  nemorum  (of  the  groves),  Con- 
volvulus sepium  (of  the  hedges),  Cassia  pumilio  (the  dwarf)  ; 
more  commonly  it  is  a  substantive  proper  name,  and  this  usu- 
ally an  old  generic  name  reduced  to  that  of  a  species.     Ex. 
Ranunculus  Flammula,  R.  Thora,  and  R.  Cymbalaria  ;  also  Lirio- 


stood  by  the  author,  and  justly  deserving  to  be  criticised."  But  this  is 
somewhat  too  absolute,  since  it  is  allowed  that  obvious  errors  in  the  con- 
struction of  names  of  Latin  or  Greek  derivation  may  be  corrected,  provided 
the  change  does  not  affect  the  initial  letter  or  syllable,  and  that  no  ancient 
names  are  to  be  disturbed. 

The  clause  that  forbids  changes  in  the  orthography  of  ancient  names, 
even  to  make  them  classical,  is  a  very  proper  one.  The  botanical  Latin 
of  Tournefort,  Linnaeus,  Jussieu,  and  their  contemporaries,  has  by  pre- 
scription rights  which  botanists  are  bound  to  respect.  Wherefore  Pyrus 
is  the  botanical  name  of  the  pear-tree,  notwithstanding  the  classical  Pints. 
So  l&vis,  as  a  specific  name  for  a  smooth  plant  (and  as  distinguished  from 
ttvis,  a  light  or  slight  one),  is  fixed  by  long  botanical  use,  although  only 
levis  is  classical ;  and  it  is  unnecessary  to  change  Ranunculus  acris  to  R.  acer. 


NOMENCLATIVE.  351 

dendron  Tulipifera,  Rhus  Toxicodendron,  Dictamnus  Fraxinella. 
These  proper  specific  names  take  a  capital  initial  letter.1  Rarely 
such  a  name  is  in  the  genitive  ;  as  Heterotheca  Chrysopsidis,  mean- 
ing a  species  of  Heterotheca  with  the  aspect  of  a  Chrysopsis. 

713.  Specific  names  should  be  of  a  single  word.     Some  few 
are  compounded,   as  purpureo-cceruleum ;  and  some  of  ancient 
origin   (once  quasi-generic)   are  of  two  words.     Ex.  Panicum 
Crusgalli,  Capsella  Bursapastoris,    Taraxacum  Dens  leonis. 

714.  A  specific  name   cannot   stand   alone.      It  is  nothing 
except  as  connected  with  the  genus  to  which  it  pertains.     A 
Japonica  by  itself  is  wholly  meaningless.     A  plant  is  named  by 
the  mention  of  its  generic  appellation  followed  by  the  specific. 

715.  Names  of  Varieties.      These  are  in  all  particulars  like 
specific  names.     Many  are  specific  names  reduced  to  a  lower 
rank.     The  varietal  name  is  written  after  the  specific,  thus : 
Ranunculus  Flammula,  var.  reptans,  and  R.  aquatilis,  var.  tricho- 
phyllus.     Varieties  of  low  grade  need  not  be  named.     They  may 
be  designated  by  numbers,  or  by  the  small  letters  of  the  Greek 
alphabet,  «,  |3,  &c.     When  the  varieties  are  marked  a  and  |3, 
the  first  is  supposed  to  be  the  type  of  the  species,  or  both  to 
be  equally  included  in  the  common  character.     But  when  the  a 
is  not  used,  the  varieties  rank  as  deviations  from  the  assumed 
type  of  the  species.     Varieties  of  cultivation,  half-breeds  or 
cross-breeds,  and  the  like,  should  have  only  vernacular  names, 
at  least  not  Latin  ones  such  as  may  be  confounded  with  true 
botanical  names. 

716.  Names  of  Hybrids  are  difficult  to  settle  upon  any  com- 
plete s}-stem.     When  of  unknown  or  uncertain  parentage,  they 
have  been  named  in  the  manner  of  species,  but  distinguished  by 
the  sign  X  prefixed.    Ex.  X  Salix  capreola.     Hybrids  of  known 
parentage  are  named  by  combining  the  names  of  the  two  pa- 
rents, thus :    S.  purpureo  X  daphnoides,  or  X  S.  purpureo-daph- 
noides,  for  a  cross  between  S.  purpurea  and  S.  daphnoides,  of 
which  the  first  supplied  the  pollen  to  fertilize  the  second.     The 
counterpart  hybrid  is  X  &  daphnoideo-purpurea. 


1  In  respect  to  the  initial  of  geographical  specific  names,  being  adjec- 
tives, such  as  Americana,  Canadensis,  Virginiana,  Europcea,  Anglica,  usage 
governs,  and  this  is  divided.  But  the  elder  DeCandolle,  who  ruled  in  all 
such  matters  in  the  preceding  generation,  always  employed  the  capital  in- 
itial, and  two  generations  of  DeCandolle  follow  the  example.  Most  English 
authors  until  recently  and  some  continental  ones  adopt  this  usage ;  and  it 
accords  with  the  genius  of  the  English  language,  in  which  we  always  write 
European,  British,  American,  &c.,  with  a  capital  initial.  Of  late  it  is  a  usual 
practice  to  write  such  geographical  specific  names  with  a  small  initial 


352  PHYTOGRAPHY. 

717.  The  Fixation  and  Precision  of  Names.     The  name  of  a 

plant  is  fixed  by  publication,  and  takes  its  date  from  the  tune 
when  it  is  thus  made  known  to  botanists. 

718.  A  genus  or  other  group  is  published  when  its  name 
and  characters  (or  the  differences  between  it  and  all  other  such 
groups)  are  printed  in  some  book,  journal,  or  other  adequate 
vehicle  of  publication,   which  is  placed  on  public  sale,  or  in 
some  equivalent  way  is  distributed  among  or  within  the  reach 
of  botanists.    A  printed  name  without  characters,  and  charac- 
ters without  name,  do  not  amount  to  publication.1 

719.  A  species  is  not  named  unless  it  has  assigned  to  it  both 
a  generic  and  a  specific  name.     It  is  not  published  until  it  is 
made  known,  by  name  and  characters  (or  by  name  along  with 
sufficient  information  as  to  its  characteristics),  in  the  manner 
aforesaid.   (718.)  Adequate  distribution,  among  botanists  and 
public  herbaria,  by  sale  or  otherwise,  of  a  collector's  or  distrib- 
utor's specimens,  accompanied  by  printed  or  autograph  tickets, 
bearing  the  date  of  the  sale  or  distribution  (that  is,  publication 
by  named  Exsiccatce  in  place  of  printed  descriptions) ,  is  held  to 
be  tantamount  to  publication.2 

720.  Characters,  references  to  date  and  place  of  publication, 
and  the  like,  belong  to  bibliography  or  particular  phytography, 
not  to  nomenclature  ;  but  proper  identification  of  names  requires 
that  the  name  of  the  author  and  the  time  and  medium  of  pub- 
lication should  be  taken  into  account.     Anterior  to  the  binomial 
nomenclature,  the  botanical  name  of  the  common  tall  Buttercup 
was  "  Ranunculus  pratensis  erectus  acris"  according  to  Bauhin, 
in  his  Pinax,  p.  179.     Under  the  new  nomenclature,  which  re- 
duced the  specific  part  of  the  plant's  name  to  one  word,  this 
became  Ranunculus  acris  in  Linnaeus,  Species  Plantarum  (ed.  1), 
p.  554 ;   and  a  brief  character  gave  its  distinctions.     In  later 
works  it  has  been  more  fully  described,  in  some  illustrated  by 
figures.     The  citation  of  these  works  arranged  in  chronological 
order  (or  in  some  order) ,  with  reference  to  volume,  page,  and 
in  some  cases  figures,  is  the  bibliography  of  the  plant.8     A  bot- 

1  Names  may  be  communicated,  in  manuscript  or  otherwise,  by  the  pro- 
pounder  to  an  author  who  may  make  them  known  by  publication ;  but  the 
date  of  the  genus  or  other  group  is  that  of  actual  publication. 

2  This  does  not  cover  all  the  conditions  of  publication,  since  it  does  not 
specify  the  characters  (and  the  same  may  be  said  of.  a  published  figure, 
with  analyses) ;  but,  on  the  other  hand,  it  conveys  to  the  competent  person 
receiving  the  same  all  this  information  and  more :  so  that  it  should  carry 
the  rights  of  true  publication  as  against  any  author  to  whom  such  names 
are  or  should  be  known.     That  is,  such  are  not  in  the  category  of  "  unpub- 
lished names,"  which  generally  ought  to  be  left  untouched. 

*  For  good  examples  of  bibliography,  see  such  detailed  works  as  De- 


NOMENCLATURE.  358 

anist,  in  referring  to  this  or  any  other  plant,  might  cite  any  work 
which  describes  it,  or  none  at  all.  Ranunculus  acris  by  itself, 
as  it  happens,  would  lead  to  no  ambiguity.  Not  so  with  many 
names.  For  the  accurate  indication  of  the  species,  it  is  generaDy 
needful,  or  highly  convenient,  to  specify  at  least  the  name  of 
the  author  who  first  published  the  adopted  appellation.  So  we 
write  Ranunculus  acris,  Linn.,  orZ.,  the  abbreviated  name  of 
Linnaeus.1  Here  we  have  the  name  of  the  plant,  and  the  bibli- 
ography reduced  to  its  initial.  To  this,  further  citation  and 
other  references  may  be  added  or  not,  as  the  particular  case 
requires.  But,  so  far  as  citation  or  reference  proceeds,  it  should 
simply  state  the  history  correctly  and  clearly. 

721.  When  a  species  is  said  to  be  of  Linnaeus  or  DeCandolle 
or  Bentham,  it  is  simply  meant  that  the  adopted  name  of  the 
plant  (consisting  of  the  generic  and  specific  parts  together)  was 
first  published  by  this  author.  Some  other  author  may  have 
named  it  differently,  and  even  earlier.  The  earlier  name  may 
have  been  discarded  because  the  specific  portion  of  it  was  un- 
tenable, either  on  account  of  preoccupation  or  for  other  valid 
reasons.  Or  the  later  author  may  have  differed  from  the  earlier 
in  his  views,  and  have  referred  the  plant  to  some  other  genus. 
As  instances  of  the  first,  Euphorbia  nemoralis,  DarL,  is  a  good 
species,  first  named  by  Darlington  in  his  Flora  Cestrica.  But 
the  name  of  Euphorbia  nemoralis  had  already  been  applied  to 
and  was  the  recognized  name  of  a  different  species  of  the  south 
of  Europe.  Whereupon,  as  the  North  American  species  had 
no  other  trivial  name,  a  new  one  had  to  be  given  to  it ;  and  it 
was  named  E.  Darlingtonii,  in  honor  of  the  discoverer  and  first 
describer.  The  common  Milkweed  of  Atlantic  North  America 
was  named  by  Linnaeus  Asckpias  Syriaca.  As  this  plant  is  not 
indigenous  to  any  part  of  the  Old  World,  and  does  not  at  all 
inhabit  Syria,  this  trivial  name  is  not  merely  faulty  but  false ; 
so  it  was  changed  by  Decaisne  into  A.  Cornuti,  in  commemora- 
tion of  an  ante-Linnaean  botanist  who  collected  it  in  Canada  and 
gave  the  first  account  and  figure  of  it.  As  an  instance  of  the 
second,  take  the  pretty  little  vernal  plant  Anemone  thalictroides, 
L.,  meaning  an  Anemone  resembling  a  Thalictrum.  When  it 
was  seen  that  the  essential  characters  were  rather  those  of  Tha- 
lictrum, the  plant  was  placed  in  the  latter  genus.  This  was  first 
done  in  Michaux's  Flora  ;  and  so  the  accepted  name  is  Thalictrum 

Candolle's  Systema  Vegetabilium,  and  Sereno  Watson's  Bibliographical 
Index  to  North  American  Botany,  in  the  Smithsonian  Miscellaneous  Col- 
lections. 

1  For  Abbreviations  of  Authors'  Names,  see  385. 


354  PHYTOGRAPHY. 

anemonoides,  Michx.,  meaning  an  Anemone-like  Thalictrum,  and 
Michaux  is  the  authority  for  this  name.  The  names  which  for 
any  reason  are  superseded  become  Synonyms*  (755.) 

722.  A  later  author  ma}T  circumscribe  a  species  or  a  genus 
differently  from  the  originator  of  the  name.  To  a  greater  or 
less  extent,  this  must  continually  happen  in  the  course  of  time. 
But  "  Ricinus  communis,  Linn."  stands  unmoved  by  the  sub- 
sequent admission  of  various  species  (known  or  unknown  to 
Linnaeus)  and  the  final  reduction  of  all  to  one  by  a  thorough 
monographer.  So  does  Silene  Gattica,  Linn.,  although  S.  quin- 
quevulnera,  Linn.,  of  the  same  date,  is  reduced  to  it.  There  is 
no  sufficient  reason  for  writing  Myosotis,  Brown,  or  Cynoglossum, 
Brown,  because  this  author  restricted  the  limits  of  these  genera ; 
nor  to  write  Gilia,  Benth.,  because  Bentham  vastly  extended 

1  The  synonymy  is  an  essential  part  of  the  bibliography  or  scientific 
history  of  a  genus  or  species.  But  synonymous  and  admitted  names  ought 
to  be  kept  distinct.  Keeping  this  principle  in  view,  —  also  the  decisively 
affirmed  doctrine  of  the  founder  of  our  nomenclature,  that  the  specific  name 
is  a  nullity  apart  from  the  generic  (so  that  only  the  combination  of  the  two 
makes  the  name  of  the  plant,  as  truly  as  the  constituent  halves  make  the 
scissors),  and  bearing  in  mind  the  fundamental  importance  and  absolute- 
ness of  the  rule  that  no  new  names  ought  to  be  m  ide  where  there  are  tena- 
ble old  ones,  —  the  student  need  not  be  misled  by  the  confusing  (however 
specious)  innovation  countenanced  by  many  zoologists  and  some  botanists, 
and  which  has  of  late  years  been  very  fully  discussed. 

The  true  rule  is  :  "  For  the  indication  of  the  name  or  names  of  any  group 
to  be  accurate  and  complete,  it  is  necessary  to  quote  the  author  who  first 
published  the  name  or  combination  of  names  in  question."  (A.  DC  )  Thus, 
Leontice  thalictroides,  Linn.,  fulfils  the  condition,  except  where  a  reference  to 
the  work  as  well  as  the  name  of  the  originator  of  the  name  is  demanded. 
Then  the  citation  would  continue,  "  Spec.  PL  312,"  and  might  be  further  ex- 
tended. In  the  Flora  of  Michaux,  this  plant  was  treated  as  distinct  from 
Leontice  in  genus ;  and  some  botanists  adopted  this  view,  while  others  of 
equal  authority  did  not.  Those  who  adopt  Michaux's  genus  name  the  plant 
Caulophyllum  thalictr -aides,  Michx. 

Now  some  naturalists  quote  for  the  species  the  author  who  originated 
the  trivial  appellation  even  when  transferred  to  another  genus.  They 
would  adopt  the  genus  Caulophyllum,  yet  write :  Caulophyllum  thalictroides, 
Linn.  Or  else  they  would  avoid  direct  falsification  of  the  facts  by  adding 
(sp.),  this  being  explained  to  mean  that  the  specific  part  of  the  name  only 
was  given  by  Linnaeus.  Then,  as  this  omits  all  mention  of  the  original  gen- 
eric part  of  the  name,  others  add  this  in  a  parenthesis,  and  write :  "  Caulo- 
phullum  thalictroides  (Linn,  sub  Leontice)  Michx.,"  or  "Caulophyllum  (Michx.) 
thalictroides,  Linn,  sub  Leontice,"  or  "Caulophyllum  (Leontice,  Linn.)  thalictroides, 
Michx."  All  such  endeavors  to  mix  synonymy  with  nomenclature  appear 
to  be  faulty  in  principle  and  unwieldy  in  practice.  In  the  most  abbreviated 
form,  they  state  that  which  is  not  true :  in  the  others,  they  impair  the  sim- 
plicity and  brevity  of  the  binomial  nomenclature.  It  is  all  but  certain  that, 
if  the  genus  Caulophyllum  had  been  published  in  the  lifetime  of  Linnaeus, 
he  would  not  have  adopted  it 


NOMENCLATURE.  355 

the  comprehension  of  this  genus.  Yet  in  their  proper  place 
such  changes  may  be  indicated  by  "pro  parte"  or  "  char,  muta- 
tis" "  excl.  sp."  and  the  like,  —  useful  qualifying  statements, 
but  no  part  of  the  name. 

723.  Exactness  requires  that  when  a  group  is  changed  from 
a  higher  to  a  lower  rank,   or  the   opposite,  the  name  of  the 
author  who  made  the  change  should  be  quoted.1     He  alone  is 
responsible  for  it.     But  this  rule  has  only  recently  been  strictly 
observed. 

724.  In  transferring  a  species  from  one  genus  to  another,  its 
specific  name  must  be  preserved  (with  alteration  of  the  gender, 
if  need  be) ,  unless  there  is  cogent  reason  to  the  contrary.     It 
must  necessarily  be  changed  when  there  is  alread}'  in  that  genus 
a  species  of  the  same  name ;  and  then  synonymous  names  of 
the  transferred  species  have  their  claim  in  order  of  date.     But 
whatever  name  is  first  employed  under  the  accepted  genus,  being 
unobjectionable,  should  hold,  even  against  an  older  unobjection- 
able one  coming  from  a  wrong  genus.     This  is  an  application 
of  the  stringent  rule  that  no  needless  names  should  be  created.* 


1  Thus,  Potentilla  Canadensis,  L.,  var.  simplex,   Torr.  $•  Gray,  and  not  of 
Michaux,  for  it  is  the  species  P.  simplex,  Michx.      Geum,  subgen.  Styliput 
Torr.  $•  Gray,  not  of  Raf.,  for  it  is  the  genus  Stylipus  of  Rafinesque,  who 
neither  made  the  subgenus  nor  approved  it.     So,  also,  for  the  genus  Labur- 
num we  write  "Laburnum,  Griseb. ; "  for  even  if  it  exactly  corresponded  with 
Cytisus  sect.  Laburnum  of  DeCandolle,  the  latter  is  not  a  group  of  equiva- 
lent rank. 

But,  as  to  genera  and  subgenera,  this  precision  should  not  be  insisted  on 
for  times  quite  anterior  to  the  recognition  of  such  rules  and  of  their  need. 
Spergularia  began  with  Persoon  as  a  subgenus  in  the  year  1805,  and  this 
date  has  been  assigned  to  the  genus,  although  it  was  taken  up  as  such  only 
in  1819  by  Presl  and  in  1824  by  Bartling. 

2  Thus,  hi  the  case  of  an  older  specific  name  being  known,  as  that  of 
Chilopsis  saligna,  Don,  recognized  as  Bignonia  linearis,  Cav.,  though  Don  ought 
to  have  adopted  the  latter  trivial  name,  yet  as  he  did  not  (and  the  rule  was 
not  then  really  in  force  as  now),  there  was  no  need  for  the  introduction 
of  a  third  name,  Chilopsis  linearis,  DC.    "  So,  again,  an  Indian  Grass  was 
first  named  and  described  by  Willdenow  as  Coix  arundinacea,  then  named 
by  Roxburgh  as   Coix  barbata,  and   entered  in   Sprengel's  Systema  with 
Willdenow's  character  as  Coix  Kcenigii.    All  these  names  were  defective 
as  referring  to  a  wrong  genus.     Brown  corrected  the  error  by  creating  the 
new  genus  Chionachne,  and  selected  Roxburgh's  specific  name  as  the  one 
most  generally  known  and  the  least  liable  to  misinterpretation ;  and  Brown's 
Chionachne  barbata  is  therefore  the  first  correct  name ;  for  which  Thwaites 
afterwards  substituted  Chionachne  Kcenigii,  an  entirely  new  and  useless  name, 
which  falls  by  the  law  of  priority.     It  should  be  well  borne  in  mind  that 
every  new  name  coined  for  an  old  plant,  without  affording  any  aid  to 
science,  is  only  an  additional  impediment."    Bentham  (Notes  on  Euphorbi- 
aceae,  in  Jour.  Linn.  Society,  xvii.  197,  198,  November,  1878).    The  following 


356  PHYTOGRAPHY. 

725.  Names  of  Snbgenera  or  of  other  sections  of  genera  are 
like  those  of  genera ;  indeed  very  many  of  them,  and  the  most 
fitting,  are  old  generic  names  which  have  been  comprehended 
in  the  genus  by  reduction.  Unlike  genera  and  higher  groups, 
however,  sections,  when  of  Greek  derivation,  may  properly  take 
the  termination  in  -oides,1  and  the  typical  section  may  bear  the 
name  of  the  genus  with  the  prefix  Eu.2  Sections  need  not  be 
named  at  all,  and  only  those  of  comparatively  high  rank  should 

is  a  farther  extract  from  the  same  protest  against  the  practice  "of  creating 
a  new  name  in  order  to  combine  an  old  specific  with  a  new  generic  one :  " 
"  In  Ferns,  the  wanton  multiplication  of  ill-defined  or  undefinable  genera, 
according  to  the  varied  fancies  of  special  botanists,  has  had  the  effect  of 
placing  the  same  species  successively  hi  several,  sometimes  seven  or  eight, 
different  genera ;  and  it  is  proposed  to  maintain  for  the  specific  appellation 
the  right  of  priority,  not  only  in  the  genus  alone  in  which  it  is  placed,  but 
in  the  whole  of  the  genera  to  which,  rightly  or  wrongly,  it  has  been  referred. 
This  has  been  carried  to  such  an  extent  as  to  give  to  the  specific  name  a 
general  substantive  aspect,  as  if  the  generic  ones  were  mere  adjuncts, — 
a  serious  encroachment  on  the  beautiful  simplicity  of  the  Linnaean  nomen- 
clature ;  and  it  is  to  be  feared  that  there  is  a  tendency  in  that  direction  in 
phaenogamic  botany.  When  a  botanist  dismembers  an  old  genus,  the  rule 
requires  that  he  should  strictly  preserve  the  old  specific  names  in  his  new 
genera ;  and,  when  he  has  wantonly  and  knowingly  neglected  this  rule,  it 
may  be  right  to  correct  him.  But  where  a  botanist  has  established  what 
he  believes  to  be  a  new  species,  and  has  therefore  given  it  a  new  name,  the 
changing  of  this  name  after  it  has  got  into  general  circulation,  because  it 
has  been  discovered  that  some  other  botanist  had  previously  published  it 
in  a  wrong  genus,  is  only  adding  a  synonym  without  any  advantage  what- 
ever, and  is  not  even  restoring  an  old  name ;  for  the  specific  adjective  is 
not  of  itself  the  name  of  a  plant.  ...  A  generic  name  is  sufficiently  in- 
dicated by  one  snbstantive ;  for  no  two  genera  in  the  vegetable  kingdom 
are  allowed  to  have  the  same  name ;  but  for  a  species  the  combination  of 
substantive  and  adjective  is  absolutely  necessary,  the  two-worded  specific 
name  is  one  and  indivisible ;  and  combining  the  substantive  of  one  with  the 
adjective  of  another  is  not  preserving  either  of  them,  but  creates  an  abso- 
lutely new  name,  which  ought  not  to  stand  unless  the  previous  ones  were 
vicious  in  themselves,  or  preoccupied,  or  referred  to  a  wrong  genus.  It  is 
probably  from  not  perceiving  the  difference  between  making  and  changing 
a  name  that  the  practice  objected  to  has  been  adopted  by  some  of  the  first 
among  recent  botanists."  Bentham,  1.  c. 

1  A  genus  could  not  properly  have  one  of  its  sections  called  by  its  own 
name  with  the  addition  of  -oides  or  -opsis,  as  Asteroides  or  Asteropsis,  for  it 
is  senseless  to  declare  that  an  Aster  resembles  an  Aster ;  but  sectional  names 
of  this  composition  may  be  excellent  for  sections  of  other  genera,  as  ex- 
pressing analogy  or  resemblance.     Latin  generic  names  used  for  sectional 
ones  properly  take  the  addition  of  -ella,  or  -ina,  or  -astrum. 

2  The  prefix  Eu  (Greek  for  much,  very,  or  true),  prefixed  to  a  generic 
name  of  Greek  origin,  is  the  proper  designation  of  the  typical  section  of 
that  genus,  meaning  the  group  which  should  bear  the  generic  name  if  such 
genus  were  divided.     The  rule  against  hybrid  names  should  in  strictness 
exclude  this  prefix  from  Latin  names,  but  it  has  not  always  done  so. 


NOMENCLATURE.  357 

have  substantive  names.  Designations,  however,  are  conven- 
ient for  lower  sections ;  and  the  name  of  a  leading  species 
may  be  used,  in  the  plural ;  as  Aster,  section  Amelli,  and  sect. 
Goncinni.  Subgenera  need  not  agree  in  gender  with  the  genus 
they  belong  to.  When  written  with  the  name  of  the  plant, 
the  subgeneric  name  is  parenthetically  inserted  between  the 
generic  and  specific  appellation.  Ex.  Pyrus  (Mains)  coronaria. 

726.  Names  of  Tribes,  Orders,  &c.     The  names  of  all  groups 
superior  to  genera  are  adjectives  plural,  and  with  few  excep- 
tions are  the  names  of  genera  lengthened  by  some  adjective 
termination.     Ex.  From  Rosa,  Rosece,  Rosacece,  Resales;  from 
Myrtus,  Myrtece,  Myrtacece,  Myrtales  ;  from  Berberis,  Berberidece  ; 
from   Tamarix,    Tamaridnece ;   from   Salix,   Salicece,  Salicinece. 
The  substantive  Plantce  being  understood,  the  groups  are  Rose- 
ous,  or  Rosaceous,  or  Rosal  plants,  &c. 

727.  Tribal  Names,  and  names  of  whatever  grade  between  gen- 
era and  orders,  are  formed  by  adding  to  the  root  of  a  generic 
name  a  final  -ece.     Ex.  Rosece,  Phaseolece,  Antirrhinece,  Oxalidece, 
&c.     Some  subtribes  take  the  name  of  the  tribe  with  the  prefix 
Ea,  as  Euphaseolece  for  that  subtribe  of  the  tribe  Phaseoleae 
which   comprises  the  representative  genus  Phaseolus.      Tribal 
names  may  take  the  same  prefix,  as  Euccesalpinece  for  the  tribe 
of  the  suborder  Csesalpinese  which  contains  the  typical  genus 
Caesalpinia. 

728.  Ordinal  Names  are  formed  in  the  same  way,  but  with  a 
preference  for  certain  terminations  which  may  denote  their  rank, 
especially  that  of  -acece,  —  as  Rosacece,  Myrtacece,  Cucurbitacece, 
meaning  Rosaceous,  Myrtaceous,  and  Cucurbitaceous  plants. 

729.  The  names  of  what  we   now   call   natural   orders,   as 
sketched  or  adopted  by  Linnaeus,  were  mostly  descriptive,  such 
as  Ensatce,  Spathacecs,  Coronarice,  Papilionacece ,  Coniferce,  Amen- 
tacece,  Umbellatce ;  but  a  few  took  their  names  from  genera,  as 
Orchidece,  Liliacece.     Jussieu,  with  whom  the  sj^stem  of  natural 
orders  properly  began,  had  no  suborders,  tribes,  or  any  such  gra- 
dation of  groups  to  deal  with.     His  one  hundred  ordinal  names 
are  some  of  them  of  the  descriptive  kind,  as  several  of  the  above, 
also  Leguminosce,  Corymbiferos,   &c.     But  the  greater  part  are 
simply  plurals  of  generic  names,  such  as  Asparagi,  Junci,  Lilia, 
Musce,  Orchides,  Lauri,  Convolvuli,  Ericce,  Acera,  Cacti.     To  a 
few  was  given  the  lengthened  termination  in  -ece,  as  Polygonece, 
Solanece,  Berberidece,  Caryophylleos ;  to  some,  the  termination  in 
-acece,  as  in    Cichoracece,    Campanulacece,   Rubiacece,   Ranuncu- 
lacece,   Malvacece,    Tiliacece,    Cucurbitacece.     Subsequent   authors 
have  necessarily  changed  all  names  which  were  plurals  of  gen- 


358  PHYTOGRAPHY. 

era ;  and  the  strongly  prevalent  tendency  has  been  to  give  the 
termination  in  -acea  to  all  such  ordinal  names,  and  to  restrict 
this  termination  to  orders.  Lindley  insisted  upon  making  this 
an  absolute  rule  even  for  names  not  formed  from  generic  appel- 
lations ;  but  this  will  not  be  adopted. 

730.  In  the  first  place,  several  large  orders  which  have  been 
known  from  the  first  by  such  characteristic  names  as  Crucifera, 
Leguminosa  (and  its  suborder  Papilionacea) ,  Gutttfera,  UmbeUi- 
fera,  Composites,  Labiates,  Cupuliferce,  and  Gonifera,  also  Palmece 
and  Graminea,  Filices,  and  even  Aroidea  and  Ficoidea,  will  retain 
these  appellations  ;  but  no  new  ones  of  the  kind  will  be  made. 

731.  Also,  names  formed  from  genera  which  do  not  well  take 
the  termination  in  -acea  may  be  allowed  as  orders  to  retain  their 
natural  form  in  -inece,  -idea,  -ariece,  and  the  like.     Ex.  Tamaris- 
cinea,  Salicinea,  Scrophularinea,  Berberidea,  Lentibulariea.     We 
may  prefer  for  the  sake  of  uniformity  to  write  Salicacea,  Berberi- 
dacea,  Lentibulariacece,  and  Scrophulariacea  (as  we  should  write 
Violacea) ,  but  this  form  cannot  be  insisted  on.     On  the  other 

hand,  a  termination  in  -acece  has  been  allowed  in  the  names  of 
certain  tribes  to  avoid  excessive  iteration  of  vowels.  Thus,  for 
the  tribe  of  which  Vernonia  is  the  leading  genus,  authors  write 
Vernoniacea,  to  avoid  Vernoniece,  which  ends  with  four  vowels. 
Spiraea  and  Staphylea  are  the  types  of  tribes,  for  which  the  names, 
if  they  followed  the  rule,  would  be  Spirceece  and  Staphyleea, 
ending  one  in  five  the  other  in  four  consecutive  vowels.  Some 
avoid  this  by  writing  Staphyleacea  and  Spiraacea.  Others  write 
Staphylea,  but  this  is  only  the  plural  of  the  generic  name. 

732.  A  few  orders  or  other  groups  took   their   names   long 
ago  from  superseded  generic  names.      Ex.    Caryophyllacea  or 
Garyophyllea,  Onagracea  or  Onagrariea,  and  Lentibulariea. 

733.  Names  of  Cohorts  are  distinguished  by  the  termination 
in  -ales.     This  was  proposed  by  Lindley,  and  is  adopted  by 

.Bentham  and  Hooker  in  the  Genera  Plantarum.  Ex.  Ranales, 
Parietales,  Malvales,  Resales,  Passijlorales,  &c.,  most  of  them 
founded  on  the  names  of  representative  genera  and  orders. 
Euphony  requires  some  to  take  other  terminations.  Ex.  Poly- 
galina,  Caryophyllina. 

734.  Names  of  Classes  and  other  great  divisions  are  plurals, 
either  adjective  or  adjective  nouns,  expressive  of  the  leading 
character.     Ex.  Polypetala,  Gamopetala,  Apetala  ;  Angiosperma 
and    Gymnosperma ;    Dicotyledones    and    Monocotyledones.      The 
names  of  the  two  great  series  or  sub-kingdoms,  following  the 
analogy  of  the  Linnaean  classes,  end  in  -ia,  and  are  Phanogamia, 
or  Phanerogamia,  and  Gryptogamia. 


GLOSSOLOGY.  359 


SECTION  II.    GLOSSOLOGY  OR  TERMINOLOGY.1 

735.  This  is  nomenclature  as  applied  to  organs  or  parts  and 
their  modifications.     The  actual  botanical  terminology  owes  its 
excellence  in  the  first  place  to  Linnaeus,  and  then  to  DeCandolle. 
The  Theorie  Elementaire  of  A.  P.  DeCandolle  (the  first  edition 
of  which  was  published  in  1813)  is  still  classical  authority,  and 
until  recently  has  received  few  additions  as  regards  terms  need 
ful  in  phaenogamous  botany. 

736.  The  fundamental  rule  is    that  each  organ  or  part  shall 
have  a  substantive  name,  and  that  modifications  of  organs  shall 
be  designated  by  adjective  terms.    These  names  or  terms  should 
be  as  precise  as  possible  :  each  object  ought  to  be  known  by  only 
one  name,  yet  synonyms  are  unavoidable ;  and  no  term  ought 
to  be  used  with  two  different  meanings.     The  word  flower,  for 
instance,  must  not  be  used  for  a  cluster  of  flowers,  however  it 
may  imitate  the  appearance  of  one,  nor  for  the  corolla  or  other 
portions  of  a  flower.    Still,  some  terms  have  to  be  used  in  two  or 
more  senses,  to  be  determined  only  by  the  connection,  or  else  as 
having  both  a  special  and  a  more  general  meaning.     Leaf  (fo- 
lium) is  a  notable  instance.     A  bract,  to  go  no  farther,  is  a  sort 
of  leaf;  and  the  imperfect  stamens  of  a  Catalpa-flower  and 
Pentstemon  are  stamens,  although  likewise  called  staminodia : 
these  are  liable  to  be  called  sometimes  by  one,  sometimes  by  the 
other  name.     But,  however  frequent  such  ambiguities  may  be  in 
morphological  treatment,  they  are  usually  avoidable  in  descriptive 
botany,  in  which  terms  are  held  to  their  more  special  or  partic- 
ular sense.     Yet  no  rule  can  absolutely  determine  whether  leaf 
or  bract,  bract  or  bractlet,  is  the  proper  term  in  many  cases. 
Moreover,  substantive   names  must  also  be  applied  to  certain 
mere  modifications  of  the  same  organ.     In  the  same  family,  a 
simple  carpel,  differently  modified  in  fruiting,  is  an  akene  in  a 
Ranunculus,  a  follicle  in  Aquilegia,  a  berry  in  Hydrastis  and 
Actsea  ;  while  in  another  family  an  additional  line  of  dehiscence 
makes  it  a  legume.     Moreover,  in  this  latter  famity  it  is  called  a 
legume  when  it  is  not  dehiscent  at  all,  and  even  when  it  becomes 
a  drupe !     Arbitrary  rules  cannot  absolutely  fix  technical  any 
more  than  ordinary  language. 

737.  Experience  and  judgment  must  determine  what  modifi- 
cations of  organs  should  be  regarded  as  a  kind,  and  bear  sub- 

1  Although  the  former  is  the  better  name,  the  latter  is  well  established  in 
use  as  an  English  word,  and  perhaps  it  need  not  be  objected  to,  inasmuch  as 
the  Latin  terminus  comes  from  the  Greek  reppa,  of  the  same  meaning. 


360  PHYTOGRAPHY. 

stantive  instead  of  merely  adjective  names.     But  the  former 
should  not  be  unnecessarily  multiplied. 

738.  The  classical  language  of  scientific  botany  being  Latin, 
all  the  organs  of  plants  and  their  principal  diversities  are  desig- 
nated by  a  Latin  or  Latinized  name.     Modern  languages  have 
also  their  own  names   and  terms.     Greatly  to  its  advantage, 
English  botanical   terminology  has   adopted   and   incorporated 
terms  from  the  Latin  and  Greek,  with  slight  changes,  not  obscur- 
ing the  identity,  thus  securing  all  their  precision,  and  rendering 
the  simple  botanical  Latin  of  descriptions  of  easy  acquisition  to 
the  English  student. 

739.  In  a  text-book  like  this,  the  principal  names  and  terms 
applied  to  organs  and  their  leading  modifications,  as  also  those 
which  relate  to  their  action  (physiological  terms) ,  or  to  our  study 
of  them  (didactic  terms,  such  as  phytography,  phyllotaxy,  glos- 
sology), are  defined  and  illustrated  in  course.     There  remain 
the  more  numerous  and  varied  characteristic  terms,  chiefly  adjec- 
tives, applicable  to  more  than  one  or  to  all  organs,  and  which 
compose  the  greater  part  of  glossology.    These,  which  DeCandolle 
arranged   systematically  with  much   elaboration,  may  best  be 
reached  by  a  glossary  or  dictionar}-,  such  as  that  at  the  end  of  this 
volume,  which  comprises  the  substantive  terms  likewise. 

740.  From   characteristic    adjective   terms   are    derived    the 
greater  number  of  specific  names  of  plants  ;  of  which,  therefore, 
the  glossary  may  elucidate  the  meaning. 

741.  Capable  as  the  existing  sj-stem  is,  it  cannot  in  single 
words   define   all   observed   forms  and  grades,  nor  well  avoid 
various  ambiguities  of  meaning.     Some  defects  of  the  first  kind 
are  remedied  by  combining  with  a  hyphen  two  congruous  terms 
to  denote  an  intermediate  state.     Ex.  ovato-lanceolatus,  or  ovate- 
lanceolate,  for  an  outline  between  the  two.     Also  a  term  may  be 
qualified  by  the  prefix  sub,  in  the  sense  of  somewhat,  as  in  sub- 
rotundus,  subcordatus  (somewhat  round  or  slightly  heart-shaped) , 
or  diminutives  (such  as  integriusculus) ,  or  superlatives  (integer- 
rimus)  or  other  strengthened  forms  (such  as  perangustus}  may 
be  employed.     Among  terms  of  more  than  one  form  of  meaning 
are  such  as  calycinus,  which  may  mean,  according  to  the  context, 
pertaining  to  the  catyx,  or  of  the  appearance  of  calyx ;  cymosus 
may  mean  in  cymes,  or  bearing  cymes,  or  in  the  manner  of  a 
cyme  ;  and  paleaceus  may  mean  provided  or  beset  with  chaff,  or 
resembling  chaff  in  texture.     Often  the  form  of  the  word  should 
distinguish  the  sense  ;  &sfoliatus,  furnished  with  leaves,  foliosus, 
with  abundance  of  leaves,  while  foliaceus  may  mean  either  bear- 
ing leaves,  or  properly  of  leaf-like  texture  or  appearance. 


DESCRIPTION.  361 

742.  Absence  of  an  organ  or  quality  may  be  expressed  by 
means  of  a  prefix  with  privative  signification,  as  indehiscent,  not 
dehiscent,    exannulate,    destitute   of   a   ring,   apetalous,   without 
petals.     But  the  Greek  privative  a  should  not  be  prefixed  to 
Latin  words,  nor  the  Latin  sub  to  terms  taken  from  the  Greek. 

743.  When  the  Latin  preposition  ob  is  prefixed  to  an  adjective 
term,  it  means  obversely ;  thus  obcordatus  is  cordate  inversed, 
that  is,  the  broader  end  with  its  notch  at  the  apex  (instead  of 
the  base)  of  the  leaf  or  other  plane  organ. 

SECTION  III.     DESCRIPTION. 

744.  Under  this  head  may  be  conveniently  comprised  all  that 
relates  to  the  form  of  the  exposition,  in  botanical  terms,  of  the 
differences  by  which  the  species  and  groups  of  plants  are  distin- 
guished and  recorded,  the  structure  exemplified,  and  the  history 
or  bibliography  indicated  in  systematic  works  or  writings.     Lin- 
naeus, in  the  Philosophia  Botanica,  treated  these  topics  under 
the  head  of  "  Adumbrationes." 

745.  Descriptions   may  be  full  and  general,   comprising   an 
account  of  all  that  is  known  of  the  structure  and  conformation 
of  a  plant  or  group,  or  rather  all  that  is  deemed  worth  recording, 
or  they  may  be  restricted  to  what  is  thought  most  important.    In 
the  former,  the  description  is  independent  of  all  relative  knowl- 
edge, or  takes  no  notice  of  relationship  to  other  plants  or  groups. 
The  latter  intends  to  portray  the  species  or  group  in  its  relations 
to  others,  and  to  indicate  the  differences  solely.     Exhaustive 
descriptions  of  the  former  kind  are  seldom  drawn  up,  but  partial 
or  supplementary  ones  are  common.     Descriptions  of  the  latter 
kind,  when  reduced  to  what  is  essential  or  differential,  are  termed 
Characters,  or  the  Character,  of  the  group  so  described.     There 
are  all  gradations  in  practice  between  characters  and  descrip- 
tions ;  but  the  distinction  should  be  maintained. 

746.  Characters  are  specific,  generic,  ordinal,  &c.     They  are 
the  differentiae,  or  marks  which  distinguish  a  group  from  any 
related  group  of  the  same  rank  with  which  it  may  properly  be 
compared.     According  to  the  occasion  and  purpose,  they  may 
specify  only  the  fewest  particulars  which  will  serve  as  a  diag- 
nosis, or  they  may  be  extended  to  all  the  known  constant  differ- 
ences between  two  or  more  related  species,  genera,  orders,  &C.1 

1  The  former  would  answer  to  what  have  been  termed  differential  char- 
acters, the  latter  to  essential  characters.  Linnaeus  divided  ( generic )  characters 
into  factitious,  essential,  and  natural;  by  the  former  denoting  any  difference 
which  may  effectively  distinguish  between  any  two  groups  brought  arti- 


362  PHYTOGBAPHT, 

What  is  now  termed  the  specific  character  was  the  specific  name 
with  Linnaeus  and  his  predecessors ;  what  we  call  the  specific, 
Linnaeus  called  the  trivial  name.  (703.) 

747.  Subordination  of  characters  and  the  avoidance  of  vain 
repetitions  require  that  as  far  as  possible  —  regard  being  had  to 
the   form   of  the  work  —  the  ordinal  character  should  contain 
only  what  is  needful  to  circumscribe  it,  and  to  exhibit  clearly 
its  morphology  ;  that  the  characters  of  tribes  or  other  divisions 
should  not  reassert  any  portion  of  the  ordinal  character,  nor 
the  generic  character  that  of  the  superior  groups  ;  and  so  of  the 
sections  and  subdivisions  of  all  grades  down  to  the  species. 
Equally  from  the  specific  character  should  be  excluded  every 
thing  which  belongs  to  the  generic,  or  is  common  to  its  rela- 
tives generally,  or  has  been  already  specified  in  the  section  or 
its  subdivisions.     So,  likewise,  of  the  varieties  under  the  spe- 
cies.    This  can  be  done  only  by  so  arranging  the  species  as  best 
to  exhibit  their  relationships,  that  is,  by  bringing  together  or 
into  proximity  those  of  greatest  resemblance  in  all  respects, 
or  in  the  more  important  respects.     What  these  are,  and  how  a 
just  subordination  of  characters  is  to  be  apprehended,  cannot 
be  taught  by  rules,  but   must  be  learned   by  experience  and 
from  the  critical  study  of  the  classical  botanical  works.     No  one 
is  competent  to  describe  new  plants  without  such  study,  and 
without  a  clear  conception  of  the   position  which  a  supposed 
new  species  should  occupy  in  its  genus,  or  a  genus  in  its  order. 

748.  Characters  of  orders,  genera,   and  of  all  intermediate 
groups,  are  drawn  almost  without  exception  from  the  organs  of 
fructification.     In  the  description,  these  parts  are  mostly  taken 
in  order,  beginning  with  the  calyx  and  ending  with  the  ovary, 
the  fruit,  seed,  embryo.    But,  as  to  the  orders,  some  writers  pre- 
fer to  preface  these  proper  characters  with  a  general  sketch  of 
those  derived  from  the  vegetation,  which,  albeit  of  less  syste- 
matic value  generally,  are  often  very  characteristic  of  particular 
families.     Rubiaceae,  for  example,  are  known  by  their  opposite 
entire  and  simple  leaves  and  intervening  stipules,  along  with  a 
few  floral  characters  ;  Sarraceniaceae,  by  tubular  or  pitcher- like 
leaves,  along  with  a  certain  combination  of  a  few  other  charac- 

ficially  together,  as  they  might  be  in  an  artificial  key,  and  as  very  'inlike 
genera  often  were  in  his  sexual  system  ;  by  the  second  meaning  the  distinc- 
tions, the  fewer  the  better,  which  will  separate  a  group  from  its  nearest 
relatives  ;  by  the  third,  all  real  marks  of  difference,  i.  e.  all  afforded  by  the 
organs  of  fructification,  which  only  were  taken  into  account  for  genera,  &c. 
Upon  the  construction  of  this  natural  character  Linnaeus  prided  himself, 
and  justly.  These  are  the  characters  in  his  Genera  Plantarum. 


CHARACTERS.  363 

ters,  and  so  on.  Where  brevity  is  aimed  at,  such  external  and 
obvious  characters,  followed  by  a  few  diagnostic  marks,  may 
practically  take  the  place  of  a  full  enumeration  of  particulars, 
many  of  which  may  be  common  to  other  orders,  though  not  in 
the  same  combination.  Generic  characters  always  commence 
with  the  calyx  or  most  external  of  the  floral  organs  and  proceed 
to  the  ovary,  thence  to  the  fruit  and  seed,  and  end  with  subsi- 
diary (but  often  no  less  diagnostic)  particulars  furnished  by  the 
vegetation  and  mode  of  growth. 

749.  Detailed  descriptions  of  species,  as  distinguished  from 
technical  characters,  commence  with  the  root,  and  proceed  in 
order  to  the  stem,  leaves  and  their  parts  or  appendages,  inflor- 
escence, bracts,  flowers,  calyx,  corolla,  stamens,  with  filament, 
anther,  and  pollen,  the  disk,  if  any,  gynoecium  and  its  parts, 
ovules ;  then  the  fruit,  seed,  albumen,   if  any,  embryo  and  its 
parts.     But  descriptions  of  this  sort  in  most  works  and  in  ordi- 
nary cases  are  partial  and  subsidiary,  comprising  only  certain 
details  supplementary  to  or  in  amplification  of  the  character  of 
the  species  or  genus.     In  condensed  works,  such  description  is 
wholly  omitted,  or  is  reduced  to  a  few  specifications  which  do  not 
readily  find  their  way  into  the  character. 

750.  Specific   characters   usually  follow  the   same   order  of 
enumeration,  from  root  to  seed,  so  far  as  the  several  organs  are 
mentioned  ;  and  in  Latin  the  phrases  are  expressed  in  the  abla- 
tive  case.     But   these  particulars  are  often  very  conveniently 
prefaced  by  statements  applying  to  the  whole  plant  rather  than 
to  any  one  organ ;  and  these  are  given  in  the  nominative,  and 
agree  with  the  name  in  gender.1 

751.  Linnaeus  required  that  neither  the  essential  character  of 
a  genus,  nor  a  specific  character  (his  nomen  specificum) ,  should 
exceed  twelve  words.     Latin  characters  take  fewer  words  than 
English.      But  this  arbitrary  rule  is  wholby  out  of  date.     Yet 
such  characters  should  be  brief  and  diagnostic  :  otherwise,  their 
advantage  is  lost,  and  the  distinction  between  them  and  descrip- 
tions  disappears.     In    monographs    and    floras,  the   desirable 
brevity,  or  such  as  the  case  admits,  is  secured  by  proper  group- 
ing under  a  subordination  of  sections,  subsections,  and  othei- 
subdivisions.2 

1  Ex.    "  NEPETA  CATARIA  :    erecta,  elata,  cano-pubescens  ;   f oliis  petio- 
Jatis,"  etc.     In  English,  these  adjectives  without  any  substantive  expressed 
will  be  seen  to  belong,  as  here,  to  "  plant "  or  "  herb  "  understood. 

2  In  the  Synoptical  Flora  of  North  America,  such  a  system  of  successive 
divisions  is  thoroughly  carried  out.    And,  if  the  specific  characters  are  by  no 
means  short,  it  is  mostly  because  nearly  all  separate  descriptive  matter  is 


364  PHYTOGRAPHY. 

752.  Punctuation.  In  proper  descriptions,  and  in  characters 
of  genera  and  of  higher  groups,  the  account  of  each  organ  forms 
a  separate  sentence  ;  and  in  Latin  the  terms  are  in  the  nomina- 
tive case,  except  subsidiary  portions,  which  are  often  thrown 
into  the  ablative.  Excepting  the  latter  part,  the  adjective  terms 
are  separated  by  commas.  A  specific  character  is  always  in  one 
sentence.  In  Latin,  its  clauses  are  mainly  in  the  ablative  ;  and 
much  diversity  prevails  as  to  the  punctuation.1  Subgeneric  and 
other  sectional  characters  are  commonly  framed  like  those  of 


dispensed  with:  consequently  various  particulars  are  added  to  the  char- 
acter which  do  not  strictly  belong  to  it.  In  Bentham's  great  Flora  Austra- 
liensis,  also  in  English,  specific  characters  are  replaced  by  a  characteristic 
synopsis  at  the  head  of  each  genus ;  and  a  terse  description  under  each 
species  completes  the  account.  Moreover,  Bentham,  in  recent  works,  sucli 
as  his  revision  of  the  Genus  Cassia,  also  that  of  the  Mimoseae,  which  have 
Lathi  characters,  writes  these  in  the  nominative  case  and  each  member  in  a 
separate  sentence,  in  the  descriptive  form,  abandoning  the  long-used  abla- 
tive form. 

1  Linnaeus  employed  only  the  comma  in  the  specific  character,  along  with 
a  subsidiary  use  of  the  colon  in  a  manner  very  unlike  its  ordinary  use  in 
punctuation,  making  it  a  point  of  less  value  than  the  comma.  Thus, 

"CHENOPODICM  ALBUM  foliis  rhomboideo-triangularibus  erosis  postice 
integris  :  sumrnis  oblongis,  racemis  erectis."  Spec.  PI.  ed.  2,  319. 

Here,  while  the  two  main  members  of  the  sentence  are  separated  by  a 
comma,  a  subsidiary  portion  of  the  first  member,  relating  to  the  uppermost 
leaves,  is  separated  by  a  colon.  Linnaeus  employed  the  colon  in  the  same 
way  in  generic  characters.  This  anomalous  usage  is  now  abandoned.  But 
most  authors  have  followed  the  Linnaean  pattern  in  distinguishing  the  prin- 
cipal members  by  commas  only,  so  that  these  become  the  only  points  in  the 
specific  character,  however  complicated  that  may  be.  Thus, 

"  RANUNCULUS  ACRIS  (Linn.  Spec.  779)  foliis  pubescentibus  subglabrisve 
palmato-partitis,  lobis  inciso-dentatis  acutis,  summis  linearibus,  caule  erecto 
plurifloro  subpubescente,  pedunculis  teretibus,  calyce  subvilloso,  carpellis 
mucrone  suberecto  terminatis."  DeCandolle,  Prodromus,  i.  36. 

This  is  the  punctuation  throughout  the  Prodromus  and  in  most  contem- 
porary systematic  works.  Its  imperfection  is  shown  in  the  above-cited  speci- 
men. The  primary  members  of  the  sentence,  which  characterize  the  leaves, 
stem,  peduncles,  calyx,  and  carpels,  are  distinguished  by  the  same  grade  of 
punctuation  which  serves  for  the  parts  of  the  first  member,  viz.  the  lobes 
of  the  leaves,  and  for  a  still  subordinate  portion,  viz.  the  form  of  the  upper- 
most lobes.  This  want  of  subordination  is  to  be  remedied  by  the  use  of 
semicolons  between  the  principal  members,  and  of  the  commas  only  for  the 
secondary  ones, —  a  punctuation  now  not  uncommon,  and  which  is  adopted 
in  the  recent  first  volume  of  the  Monographiae  Phanerogamarum  of  the  De- 
Candolles,  which  supplements  the  Prodromus.  The  portion  of  that  volume 
contributed  by  Dr.  Masters  better  exemplifies  this  than  does  the  rest  of  the 
volume.  For  the  latter  sacrifices  the  advantage  of  the  change  by  the  inser- 
tion of  commas  between  each  adjective  of  a  continuous  ablative  phrase  (as, 
"  Smilax  laurifolia;  limbis  foliorum  oblongis  vel  ovato-oblongis,  coriaceis, 
3-5-nerviis,  subtus  pallidioribus,"  etc.),  where  they  are  generally  deemed 


PUNCTUATION,   SYNONYMY.  365 

genera.  Or  the  members  may  be  united  in  one  sentence,  but  in 
that  case  the  principal  ones  are  best  separated  by  colons. 

753.  Should  a  point  intervene  between  the  specific  name  and 
that  of  the  author  cited?    The  practice  varies.    But,  if  the  name 
is  Latin,  the  comma  is  superfluous  ;  for  the  abbreviated  name  of 
the  author  is  supposed  to  be  in  the  genitive,  and  to  read  thus : 
Ranunculus  repens  Linncei.     Still,  since  when  the  author's  name 
is  cited  in  full  it  is  never  written  in  the  genitive,  and  since  in 
English  the  comma  is  normall}r  required,  it  seems  on  the  whole 
proper  to  insert  it. 

754.  In  citations,  the  classical  practice  is  to  separate  the  refer- 
ences from  each  other  and  from  the  name  by  periods ;   thus, 
'•'•Anemone  cylindrica,  Gray,  Ann.  N.  Y.  Lye.  3.  221.     Torr.  & 
Gray,  Fl.  1.  113, "*  &c.      It  is  becoming  equally  customary  to 
separate  the  several  citations  by  semicolons,  thus  bringing  all  the 
references  under  one  name  into  one  sentence.    The  bibliography 
of  a  species  or  group  of  species  which  a  describer  or  other  author 
has  to  refer  to  (with  more  or  less  fulness,  according  to  the  form 
of  his  work)  is  to  be  sought  partly  under  the  admitted  name, 
and  partly  in  the 

755.  Synonymy.     This  includes  all  other  than  the  admitted 
names.    Ex.  Hydropeltis  of  Michaux  is  a  S}'nonym  of  Brasenia  of 
Schreber,  the  latter  being  the  earlier  published  name.     Nectris 
of  Schreber  is  a  synonym  of  Cabomba  of  Aublet,  the  latter  hav- 
ing priority.      Thalictrum  anemonoides  of  Michaux  has  for  syno- 
nyms Anemone  thalictroides  of  Linnaeus  and  of  many  subsequent 
authors  who  followed  him  in  referring  this  ambiguous  plant  to 
Anemone  (721) ;  and  also  Syndesmon  thalictroides  of  Hoffmannsegg 
and  Anemonella  thalictroides  of  Spach,  who  proposed  to  consider 

superfluous.  The  preferable  punctuation  of  the  character  above-quoted 
from  the  Prodronnis  would  be 

RANUNCULUS  ACRIS  (Linn.):  foliis  pubescentibus  subglabrisve  palmato- 
partitis,  lobis  inciso-dentatis  acutis,  summis  linearibus ;  caule  erecto  pluri- 
floro  subpubescente ;  pedunculis  teretibus ;  calyce  subvilloso ;  carpellis 
mucrone  suberecto  terminatis. 

The  advantages  of  this  style  of  punctuation  will  more  and  more  appear, 
when  applied  to  less  simple  cases.  Commas  between  the  ablative  adjectives 
are  superfluous  and  confusing. 

In  English  characters,  commas  are  required  between  the  adjectives  which 
follow  the  noun.  Rightly  to  express  the  subordination  of  characters,  the  plan 
adopted  in  the  Synoptical  Flora  of  North  America  is  recommended ;  that  is, 
with  colons  separating  the  principal  members,  semicolons  for  subordinate 
and  dependent  ones,  and  commas  between  the  adjectives  of  the  same  noun. 

1  See  Watson's  Bibliographical  Index  to  North  American  Botany  (where 
this  style  is  adopted)  for  a  general  model  for  the  arrangement  of  synonym/ 
and  citations. 


366  PHYTOGBAPHY. 

it  an  intermediate  genus  between  Thalictrum  and  Anemone.  In 
systematic  works,  the  specific  character  immediately  follows  the 
name,  and  generally  forms  a  part  of  the  same  sentence ;  and  is 
followed  first  by  citations  of  authors  who  have  adopted  the  name, 
and  then  by  the  synonymy,  or  as  much  of  it  as  the  plan  of  the 
work  calls  for.  The  synonymous  names  and  the  references 
under  them  should  be  cited  in  the  order  of  their  publication. 
But,  to  economize  space,  all  the  authorities  for  the  same  name 
are  brought  together  into  one  sentence,  and  arranged  according 
to  their  date.  Also,  where  the  synonymy  is  not  elaborately 
displayed,  the  various  synonyms  of  the  same  generic  name  are 
usually  placed  in  consecutive  order. 

756.  Iconography.     The  leading  and  most  essential  citation  is 
that  of  the  author  by  whom  and  the  work  in  which  a  plant  is 
named  and  described,  and  also  the  work  in  which  it  is  best  char- 
acterized.    Among  the  characterizations,  published  figures  hold 
a  prominent  place.     The  citation  of  these  is  an  important  part 
of  the  synonymy.     The  best  botanical  plates  are  those  which 
give  detailed  analyses  of  the  parts  of  the  flower,  fruit,  and  seed, 
displaying  their  structure. 

757.  Habitat  and  Station  are  recorded  in  a  sentence  or  para- 
graph following  the  name,  character,  and  synonymy  of  a  species. 
The  habitation  is  the  place,  district,  or  region  at  or  within  which 
the  plant  is  known  to  be  indigenous,  or  to  grow  spontaneously. 
The  complete  habitat  is  the  geographical  range.     The  station  is 
the  situation  it  affects,  whether  in  water,  in  marshes,  on  shores, 
on  hills  or  mountains,  in  forests,  on  open  plains,  &c. 

758.  Discoverer,  &c.     To  the  habitat  and  station  of  newly 
discovered,  rare,  or  local  plants  should  be  appended  the  name 
of  the  discoverer  or  the  collectors  by  whom  the  species  has  be- 
come known  to  science,  at  least  when  the  plant  is  first  published. 
Date  of  discovery  should  also  then  be  indicated. 

759.  Time  of  Blossoming  should  be  recorded,  either  the  month 
or  the  season,  to  which  may  be  added  that  of  the  maturity  of  the 
fruit.     When  the  month  or  season  is  mentioned  without  farther 
explanation,  flowering-time  is  intended.      In  a  flora,  this  may 
sometimes  be  indicated   under  the   genus  for  all  the  species. 
In  the  flora  of  an  extensive  region,  and  in  respect  to  species  of 
considerable  range  in  latitude  or  longitude,  the  time  of  flowering 
differs  so  widely  at  the  extremes  of  the  geographical  range  that 
it  cannot  well  be  specified  except  in  general  terms,  as  spring, 
summer,  autumn,  &c. 

760.  Etymology  of  Names.     When  a  new  generic  name  is  pub- 
lished, its  origin  and  meaning  should  always  be  given,  if  the 


ACCENTUATION,   ABBBEVIATION8.  367 

nature  of  the  publication  will  allow  it.  So  likewise  of  species, 
except  where  the  source  or  signification  of  the  name  is  mani- 
fest. This  is  commonly  the  case  as  respects  most  characteristic 
specific  names,  and  also  those  drawn  from  station,  habitat,  and 
the  like. 

761.  Accentuation  of  Names.     The  pronunciation  of  botanical 
names  is  settled  by  the  rules  of  Latin  prosody.     All  that  is 
usually  attempted  in  those  botanical  works  which  take  this  into 
account  is  to  mark  the  syllable  upon  which  the  principal  accent 
falls.     This  in  words  of  two  syllables  is  always  the  first ;  in 
words  of  three  or  more  syllables,  either  the  penult  (the  last  sylla- 
ble but  one)  or  the  antepenult  (next  preceding  syllable) .    When 
the  penult  is  a  long  syllable,  it  takes  the  accent ;  when  short,  this 
recedes  to  the  antepenult.     The  accentuation  may  accordingly 
be  sufficiently  indicated  by  marking  the  quantity  of  the  penult, 
either  long  as  in  Erica,  or  short  as  in  Arbutus  and  Gladiolus.    Or 
else  the  accent  may  be  marked  by  a  proper  sign,  as  Erica,  Arbutus, 
Gladiolus.     An  endeavor  has  been  made  to  represent  the  longer 
sound  of  the  vowel  by  the  grave  accent-mark,  as  Erica,  and  the 
short  by  the  acute,  as  Gladiolus.     But  this  plan  is  encumbered 
with  practical  difficulties. 

762.  Abbreviations  are  required,  both  of  the  name  of  the  au- 
thor, when  of  more  than  one  or  two  syllables,  and  of  the  titles 
of  the  works  cited.     There  are  also  the  customary  abbreviations 
in  the  citation  of  volume,  page,  plate,  &c.,  in  which  there  is 
nothing  peculiar  to  botany. 

763.  The  simple  rule  for  the  abbreviation  of  an  author's  name 
is  to  abridge  it  of  all  but  the  first  syllable  and  the  first  letter  of  the 
following  one  (as  Lam.  for  Lamarck,  Hook,  for  Hooker) ,  or  the 
first  two  letters  following  the  vowel  when  both  are  consonants 
(as  Linn,  for  Linnaeus,  Juss.  for  Jussieu,  Rich,  for  Richard) . 
Sometimes  more  of  the  name  must  be  given,  in  order  to  distin- 
guish those  beginning  with  the  same  syllable.    So  we  write  Mchx. 
to  prevent  confusion  of  the  name  Michaux  with  that  of  Micheli, 
which,  being  the  earlier,  claims  the  abbreviation   Mch.,  and 
Bertol.  to  distinguish   Bertoloni   from   Bertero.      Sometimes  a 
much-used  name  of  one  syllable  is  abbreviated,  as  Br.  (or  R. 
Br.)  for  Robert  Brown.     Initials  or  abbreviations  of  the  bap- 
tismal name  are  needed  to  distinguish  botanists  of  the  same 
name ;    as  P.  Browne  in  distinction  from  Robert  Brown,  Ach. 
Rich.,  Adr.  Juss.,  Alph.  DO.,  to  distinguish  the  younger  from  the 
older  Richard,  Jussieu,  and  DeCandolle.     Or,  where  father  and 
son,  the  abbreviation  for  the  latter  may  be  Juss.jiL,  Hook.jil., 
or  Hook.  /.,  &c.     Certain,  but  very  few,  well-known  and  eminent 


368  PHYTOGRAPHY. 

names  are  abbreviated  to  a  sign ;  as  L.  for  Linnaeus,  DC.  for 
DeCandolle,  HBK.  for  Humboldt,  Bonpland,  and  Kunth,  the 
latter  too  long  after  ordinary  abbreviation.1  Care  should  be 
taken  to  affix  the  period  by  which  abbreviations  may  be  dis- 
tinguished from  full  names,  such  as  Don,  Ker,  Blytt. 

764.  Abbreviations  of  titles  of  works  follow  the  same  rules  as 
those  of  names,  or  at  least  are  in  no  wise  peculiar  in  botany. 

765.  Abbreviations  of  the  names  of  organs  follow  the  same 
rule :    Col.   for  calyx,    Cor.  for  corolla,    Stam.  for  stamen   or 
stamina,  Pist.  for  pistillum  or  pistil,  Fr.  for  fructus  or  fruit, 
Per,  for  pericarpium  or  pericarp,  Sem.  for  semen  or  seed,  are  the 
most  common.     Hab.  for  habitat  or  geographical  station,  Herb. 
for  herbarium,   Gen.  for  genus,  Sp.  or  Spec,  for  species,   Var.  for 
variety,  and  the   like,  every  one  will  understand.      But   some 
abbreviations  which  are  common  in  botanical  writings,  at  least 
those  in  Latin,  may  need  explanation  to  the  elementary  student. 
A  list  of  abbreviations  is  appended.     See  p.  390. 

766.  Signs.     Under  this  head  might  be  ranked  such  abbrevia- 
tions as  v.  v.  for  vidi  vivam,  v.  s.  for  vidi  siccam,  to  note  that  the 
writer  has  seen  the  plant,  either  alive  or  in  a  dried  specimen';  or, 
more  particularly,  v.  s.  s.,  when  it  is  a  spontaneous  specimen 
that  has  been  examined  in  a  dried  state,  and  v.  s.  c.,  when  it  was 
a  cultivated  specimen ;  v.  v.  c.,  when  the  living  plant  was  seen 
in  a  garden  only,  and  v.  v.  s.,  when  the  spontaneously  growing 
plant  was  seen  alive.     There  are  also  proper  signs,  of  which  the 
most  common  are  those  which  indicate  the  sexes  of  blossoms, 
the  duration  of  a 'plant,  and  the  like.    Also  the  interrogation 
point  (?)  used  to  express  doubt ;  the  exclamation  point  (!)  to 
indicate  the  certainty  that  is  given  by  the  actual  sight  of  an 
authentic  original  specimen.     See  p.  391. 

767.  The  marks  used  to  indicate  the  subordination  of  sections 
under  a  genus,  or  in  the  synoptical  arrangement  of  genera,  and 
the  like,  are  not  settled  by  any  fixed  rule.    An  approved  ar- 
rangement is  to  employ  the  following  marks  in  the  given  order, 
§  *   -t-   -H-   =.     The  first  one,  for  sections  of  the  highest  order, 
takes  numerals  after  the  sign.      Ex.  §  1,  and  so  on.     When 

1  As  Alph.  DeCandolle  remarks,  the  proper  abbreviation  of  the  narn<<  he 
bears  is  Ccmd.  But  the  form  DC.  was  very  early  adopted  by  the  first  of 
the  illustrious  name,  and  has  been  continued  for  almost  three  quarters  of  a 
century.  Alphonse  DeCandolle  would  prefer  to  write  it  D.C.,  but  has  not 
adopted  that  mode,  nor  should  we ;  for  DC.  and  HBK.  are  convenient  ab- 
breviations reduced  to  signs.  But  such  forms  should  not  be  increased.  For 
ordinary  names  they  would  be  unintelligible. 

Names  which  are  not  too  long,  and  of  which  an  abbreviation  by  the  ordi- 
nary rule  is  insufficient,  such  as  Decaisne,  should  rather  be  written  in  full- 


SIGNS,   ETC.  369 

such  sections  are  followed  by  a  substantive  name,  they  are 
equivalent  to  subgenera.1  Ex.  Phacelia,  Juss.,  §  1.  Euphacelia, 
i.  e.  the  true  or  typical  Phacelia ;  §  2.  Cosmanthus,  Gray,  &c. 
Sections  next  in  rank  to  these  are  marked  with  asterisks,  *  for 
the  first,  *  *  for  the  second,  *  *  *  for  the  third  one  of  the  same 
rank.  Divisions  of  these  have  the  -t-  prefixed  ;  and  so  on  in  the 
same  way.  Still  farther  subdivisions  may  be  marked  by  the 
small  letters  of  the  alphabet  consecutively,  a,  b,  c.  When  capital 
letters  are  used  for  division  marks,  it  is  mostly  for  those  of  a 
high  grade. 

768.  Floras,  Monographs,  &c.     A  systematic  work  describing 
in  proper  order  the  plants  of  a  country  or  district  is  generally 
called  a  Flora.     A  Flora  of  a  small  district  takes  the  diminutive 
name  of  a  Florula.     A  universal  work  of  the  kind  when  it  ex- 
tends to  the  species  is  a  System,  Sy  sterna  Vegetabilium  or  Sy  sterna 
Regni    Vegetabilis.     The   latest  completed  Systema  Vegetabilium 
is   that  of  Sprengel   (1825-1828),  in  five  octavo  volumes,  on 
a  very  condensed  plan.     A  compendious  Flora  or  S}Tstema  is 
often  termed  a  Prodromus,  literally  meaning  a  forerunner  or 
preliminary  work.     But,  as  even  this  is  more  than  most  bot- 
anists are  able  to  complete,  the  name  of  Prodromus  is   now 
applied  to  works  which  are  not  intended  to  precede  fuller  ones 
by  the  same  author.     The  principal  work  of  this  kind  is  the  Pro- 
dromus Syst.  Nat.  Regni  Vegetabilis,  commenced  by  DeCandolle 
in  the  year  1824,  continued  by  his  son  Alphonse   DeCandolle 
(aided  by  various  botanists)  to  its  close  in  1873,  down  to  vol. 
xvii.,  or  essentially  twenty  very  compact  octavo  volumes,  these 
carrying  the  work  only  through  the  great  class  of  Dicotyledones. 
But  the  publication  of  the  monocotyledonous  orders  has  com- 
menced in  a  series  of  Monographs  (Monographic  Phanerogam- 
arum)  .     A  Monograph  is  a  systematic  account  of  all  the  species 
of  a  genus,  order,  or  other  detached  group. 

769.  Specimens  of  botanical  characters  and  descriptions,  cita- 
tions, &c.,  illustrating  this  chapter,  might  be  given  here.     But, 
for  those  in  Latin,  the  classical  works  of  DeCandolle  and  others, 
and   for  the  genera  those  of  Jussieu,  Endlicher,  Bentham  and 
Hooker,  may  be  taken  as  models.     In  English,  those  of  the 
latter   authors,   and   in   the    United    States    the    better-known 
writings  of  the  present  author,  especially  the  later  ones,  may 
be  referred   to. 

1  DeCandolle  in  the  Prodromus  employed  the  word  Sect.  (Sectio)  for 
what  answers  to  subgenus  or  at  least  to  the  highest  grade  of  sections ;  then 
§  1,  §  2,  &c.,  for  the  next  grade  below  subgenus  ;  and  then  the  asterisk,  and 
other  marks. 

If 


370  PHYTOGBAPHY. 


SECTION  IV.     SPECIMENS  ;   DIRECTIONS  FOR  THEIR  EXAMINA- 
TION, PRESERVATION,  &c. 

770.  Implements.     Those   necessary  for  the  examination  of 
phsenogamous  plants,  Ferns,  and  the  like,  are  a  simple  pocket 
lens,  a  simple  dissecting  microscope ;  also  a  sharp  thin -b laded 
knife  and  some  needles  of  various  fineness,  mounted  in  han- 
dles, for  dissection. 

771.  For  a  single  hand  lens,  one  magnifying  only  from  four  to 
six  diameters  is  the  most  useful.    A  doublet,  or  a  parabolic  lens 
of  Tolles,  of  about  an  inch  focus,  is  better,  but  much  more  expen- 
sive.   The  simple  stage-microscope  for  dissection  need  have  only 
two  lenses   (doublets  or  otherwise)  with  large  field  and  good 
definition,  one  of  an  inch  and  the  other  of  about  half  inch  focal 
distance ;  and  a  glass  stage  of  at  least  an  inch  and  a  half  in 
diameter.     A  compound   microscope   is  useful   for   all   minute 
investigation,  and  is  essential  in  the  study  of  vegetable  anatomy 
and  of  all  lower  cryptogamic  botany. 

772.  For  making  thin  slices,  a  razor  is  the  best  knife  ;  for  dis- 
section on  the  stage  of  the  simple  microscope,  beside  needles, 
small  scalpels  or  some  of  the  cutting  instruments  used  by  ocu- 
lists are  very  convenient.     But  an  expert  hand  is  able  to  do 
almost  every  thing  with  a  common  knife  or  scalpel  and  a  pair 
of  mounted  needles.     Slender  forceps  are  almost  indispensable  : 
those  made  for  the  use  of  dentists  are  the  best. 

773.  Analysis.     In  the  examination  of  an  unknown  plant  with 
a  view  to  its  determination,  its  whole  structure  should  be  made 
out,  so  far  as  the  materials  allow,  before  a  step  is  taken  to 
ascertain  its  name  and  place  in  the  system.     In  respect  to  the 
stem,  its  duration  and  consistence  and  its  internal  structure, 
whether  exogenous  or  endogenous,  are  to  be  noted.     As  to  the 
foliage,  the  venation  and  the  phyllotaxy,  also  the  presence  or 
absence  of  stipules,  are  most  important.      The  anthotaxy  or 
inflorescence  is  to  be  examined  and  referred  to  its  proper  t}-pe. 
In  the  flower,  the  numerical  plan  and  symmetry,  its  ground-plan 
and  the  nature  of  the  deviations  from  the  general  or  the  famil}- 
type,  are  to  be  considered ;  also  the  aestivation  or  arrangement 
of  the  parts  in  the  bud,  the  character  and  extent  of  coalescence 
and  adnation  ;  the  manner  in  which  the  anther  is  borne  upon  the 
filament,  and  its  place  and  mode  of  dehiscence,  &c.     Note  also 
whether,  when  the  blossom  is  hermaphrodite,  the  anthers  and  the 
stigmas  mature  at  the  same  or  at  different  periods.    The  placen- 
tation  and  the  character  and  position  of  the  ovules  should  be 


SPECIMENS.  371 

determined.  Two  sections  of  the  flower  should  be  made  :  one  of 
them  vertical  and  directly  through  the  centre,  in  the  manner  of 
Fig.  336-341,  —  this  will  display  the  adnation,  insertion,  &c., 
of  all  the  parts  ;  the  other  transverse  and  through  the  middle  of 
the  ovary,  also  above  the  ovary  when  this  is  inferior,  and  if  pos- 
sible in  the  unopened  but  full-grown  flower-bud ;  this,  among 
other  things,  will  bring  to  view  the  aestivation.  (Fig.  351, 386, 398, 
&c.)  Not  rarely  fruit  and  seeds  are  to  be  had  at  the  same  time, 
or  upon  the  same  specimen,  and  these  are  equally  to  be  investi- 
gated. In  fresh  seeds,  even  those  of  minute  size,  the  embryo 
may  almost  always  be  extracted  or  brought  to  view  under  the 
microscope,  either  by  tearing  away  the  seed-coat  with  needles  or 
by  sections  with  a  keen  knife.  When  hard  and  dry,  they  have 
only  to  be  soaked  or  slightly  boiled. 

774.  Diagrams  and  also  sketches  of  the  parts  should  be  made, 
such  as  those  referred  to  in  the  foregoing  paragraph.      Such 
diagrams  can  be  drawn  by  any  one  with  a  little  practice ;  and 
they  may  be  made  to  express  the  whole  floral  structure,  even  to 
the  coalescence  and  adnation.1    But  in  the  process  of  determina- 
tion the  student  should  beware  of  trusting  wholly  to  his  diagrams 
and  sketches  without  direct  verification. 

775.  Dried  specimens,  when  well   prepared   and   in  sufficient 
abundance,  in  the  hands  of  a  skilled  botanist  are  in  most  cases 
but  little  inferior  to  fresh  ones.     When  needed,  flowers,  or  clus- 
ters of  blossoms,  or  fruits  may  be  detached  and  prepared  for 
examination  and  dissection  by  somewhat  prolonged  soaking  in 
warm  water  or  by  a  short  immersion  in  boiling  water.     This  re- 
stores flower-buds  and  small  flowers  and  fruits,  or  their  parts,  to 
a  condition  not  essentially  unlike  the  living  state.    Consequently, 
the  Herbarium  or  Hortus  siccus  of  the  botanist  is  to  him  more 
essential  than  the  botanical  garden,  important  as  that  may  be. 

776.  Herborizing.2     The  collector's  outfit  will  essentially  con- 
sist of  a  Vasculum  or  botanical  box,  a  Portfolio,  a  Trowel,  a  pocket 
Lens,  and  a  small  but  stoutly  covered  Note-book.     Some  use  a1 
portfolio  only,  others  the  botanical  box  ;  but  on  a  long  excursion 
it  is  well  to  have  both.     The  former  is  preferable  in  most  cases, 
except  when  specimens  are  collected  for  the  immediate  use  of  a 

1  See  Eichler's  Bliithendiagramme  (Leipsic,  1872,  1878),  an  admirable 
work,  which  may  serve  as  a  model. 

2  These  articles,  from  paragraph  776  to  802  inclusive,  were  obligingly  pre- 
pared, at  the  author's  request,  by  LYMAN  H.  HOYSRADT,  of  Pine  Plains, 
New  York.     They  form  an  abstract  or  a  new  edition  of  a  series  of  notes  on 
the  subject  which  were  published  in  the  Bulletin  of  the  Torrey  Botanical 
Club, in  the  year  1878. 


372  PHYTOGRAPHY. 

class.  When  well  stocked  with  paper,  it  is  of  almost  unlimited 
capacity ;  and  most  plants  of  delicate  texture  (as  many  of  the 
smaller  aquatics,  and  those  with  fugacious  or  delicate  corollas) 
need  to  be  consigned  directly  to  the  paper  in  which  they  are  to 
be  pressed,  and  to  be  kept  meanwhile  under  some  pressure. 

777.  The    Vasculum  is  very  useful  for  holding  plants  that  are 
to  be  examined  fresh,  and  for  thick  roots,  large  fruits,  &c.     It 
is  made  of  tin,  and  should  be  of  oval-C3Tlindrical  shape,  about  17 
inches  long  and  4  by  6  inches  wide,  and  provided  with  a  light 
strap  to  throw  over  the  shoulder.      The  lid  opens  nearly  the 
whole  length  of  one  of  the  flat  sides  (15  by  4£  inches,  with  one 
fourth  inch  lap) ,  is  made  to  fit  as  close  as  possible,  and  fastens 
by  a  simple  spring  catch.    When  no  portfolio  is  used,  a  larger  box 
may  be  required.     Plants  may  be  kept  fresh  in  such  a  box  for 
many  days.    For  a  several-days  excursion,  when  it  is  desirable  to 
bring  home  a  large  number  of  fresh  plants,  a  tin  chest,  made  some- 
what after  the  pattern  of  an  old-fashioned  trunk,  will  be  found 
very  convenient.     It  should  be  about  21  inches  long,  10  inches 
wide,  and  10  inches  high  to  the  top  of  the  convex  and  hinged 
lid,  which  forms  the  whole  top,  and  to  which  a  handle  is  fitted. 

778.  A  good  form  of  Portfolio  is  made  of  two  pieces  of  binder's 
board  covered  with  enamel  cloth,  and  fastening  together  with  two 
long  straps  with  buckles.     Handles  similar  to  those  on  a  carpet- 
bag may  be  attached  for  carrying.     The  usual  size  of  portfolio 
is  18  by  12  inches,  but   16£  by  ll£  inches  may  be  better,  as 
there  would  then  be  little  danger  of  making  specimens  of  too 
great  length  for  the  herbarium.     (784.)    Or  the  back  may  be  of 
soft  leather,  an  inch  or  so  in  width,  and  a  light  strap  and  buckle 
at  the  front  edge  and  at  each  end.    The  portfolio  should  contain 
a  good  quantity  of  folded  sheets  of  thin  unsized  paper,  similar 
to  grocer's  tea-paper,  and  of  a  size  only  a  little  smaller  than  the 
sides  of  the  portfolio.     Very  thin  manilla  paper,  or  what  is  so 
called,  is  excellent  for  this  purpose,  being  sufficiently  bibulous 
and  rather  strong. 

779.  The  specimens  as  soon  as  gathered  should  be  laid  neatly 
in  these  folded  sheets  (called  specimen  sheets} ,  and  kept  under  a 
moderate  pressure  in  the  portfolio.     The  sheets  with  the  spec- 
imens are  afterwards  transferred  to  the  home   press,  but  the 
specimens  should  be  left  continuously  in  their  sheets  through  all 
the  changing  of  driers,  until  cured.     Indeed,  the  specimens  may 
well  remain  in  the  sheets  after  drying,  until  wanted  for  mounting 
or  for  exchanging.     For  fine  specimens,  the  use  of  this  specimen 
paper  is  very  important.     Many  plants  are  so  extremely  delicate 
and  sensitive  that  they  will  not  bear  the  least  handling  without 


HEBBORIZATION.  373 

curling  and  shrivelling,  unless  thus  enclosed :  also  without  these 
sheets  much  time  is  lost  in  transferring  small  specimens  one  by 
one  from  one  driei  to  another  in  the  drying  process. 

780.  For  digging  up  roots,  bulbs,  &c.,  a  small   and   sharp 
pointed  triangular  Trowel  or  stout  knife  will  answer.     One  of 
the  best  "diggers"  is  made  from  a  large  file.      Let  a  black- 
smith bend  the  lower  half  of  the  blade  to  a  gentle  curve,  so  that 
the  point  will  be  about  an  inch  out  of  the  true  line.     Grind  off 
the   teeth   and   re-temper  the   blade.      The   total   length  with 
handle,  which  is  over  one  third,  should  be  about  twelve  inches. 
A  leather  case  may  be  made  for  convenience  of  carriage.     The 
advantages  of  this  strong  tool  are  many.1 

781.  A  Note-book  should  be  carried  upon  every  excursion,  in 
which  the  station  of  rare  plants,  dates,  colors,  and  various  par- 
ticulars which   cannot   be   learned    from   the    specimens,   may 
be  recorded  on  the   spot,  instead  of  being  left  to   uncertain 
memory. 

782.  For  most  plants,  the  best  time  for  collecting  flowering 
specimens  is  in  the  morning,  soon  after  the  dew  has  disappeared. 
Vespertine  flowers  have  to  be  secured  earlier,  or  at  nightfall. 

783.  Care  should  be  taken  to  have  the  specimen  of  the  proper 
size,  neither  too  small  nor  too  large,  and  to  comprise  all  that  is 
necessary  for  complete  botanical  illustration,  —  flowers,   fruit, 
and   leaves,   both  cauline  and   radical  when   possible.      Inex- 
perienced botanists   suppose  that  a  small  sprig,  containing   a 
flower  or  two  with  a  few  leaves,  will  answer  all  purposes  as  a 
botanical  specimen ;  but  later  he  comes  to  know  better,  and  also 
learns  that  the  flower  is  only  one  of  the  component  parts  of  a 
specimen,  and  not  always  the  most  important  one.     In  various 
genera  and  orders,  the  fruit  is  the  most  distinguishing  character- 
istic, as  with  the  Potamogetons,  the  Cruciferse,  the  Umbelliferae, 
and  the  Cyperaceae.     With  many  plants  the  radical-leaves,  with 
others  the  character  of  the  subterranean  stem,  whether  a  rootstock, 
tuber,  conn,  or  bulb,  or  of  the  root  itself,  whether  annual,  bien- 
nial, or  perennial,  becomes  important.     Consequently,  all  the 
organs  have  their  value  in  an  herbarium  specimen,  and  each  and 
all  should  receive  due  consideration  from  the  botanist  when  col- 
lecting.     Specimens  may  be  often   secured   that  exhibit  both 

1  [There  is  an  English  herborizing  trowel  of  excellent  quality,  with  blade 
six  or  eight  inches  long,  less  than  two  inches  wide,  the  sides  slightly  in- 
curved, the  stout  shank  an  inch  and  a  quarter  wide,  and  one  sixth  of  an 
inch  thick :  this  forms  the  whole  back  of  the  handle,  the  front  of  which  is 
a  piece  of  lignum  vita;  riveted  fast  to  the  steel.  It  is  nearly  impossible  to 
break  it] 


374  PHYTOGRAPHY. 

flowers  and  fruit  in  the  same  plant,  or  fruit  may  be  frequently 
obtained  from  more  advanced  plants  at  the  same  time.  If  not, 
fruit  must  be  collected  later,  as  in  case  of  shrubs  and  trees,  of 
which  generally  only  a  branchlet  with  flowers,  or  with  flowers 
and  leaves,  can  be  gathered  first.  But  subsequently  the  fruit 
and  mature  leaves,  should  always  be  taken,  if  practicable,  from 
the  same  individual  as  the  flowers.  Of  dioecious  shrubs  or 
trees,  like  the  Willows,  each  species  should  be  represented  by 
four  pieces :  first,  the  sterile  and  fertile  catkins  will  have  to  be 
obtained,  and  the  respective  individuals  marked,  so  that  later 
corresponding  twigs  with  mature  leaves,  stipules,  and  fruit  may 
be  gathered,  and  the  specimens  rightly  matched.  , 

784.  A  specimen  should  be  so  arranged  as  to  be  no  larger 
when  pressed  than  can  be  neatly  mounted  on  the  herbarium 
paper.     A  slender  plant  not  over  three  feet  in  height  should 
generally  be  preserved  entire,  root  and  all.     This  can  be  done 
by  bending  or  partially  breaking  it  at  one,  two,  or  three  places, 
and  doubling  so  that  the  sections  will  not  rest  upon  each  other 
in  drying.     If  broken  twice,  it  may  be  neatly  arranged  in  the  N 
form  when  put  in  portfolio.     Very  large  herbaceous  plants  will 
have  to  be  divided  and  the  parts  preserved  separately,  or,  better, 
take  a  suitable  portion  of  the  upper  stem,  having  leaves,  flowers, 
and  fruit,  and  a  convenient  part  of  the  lower  stem  containing 
radical  leaves  and  with  it  sufficient  root  to  show  whether  the 
plant  is  an  annual,  biennial,  or  perennial.     Thick  stems,  roots, 
tubers,  bulbs,  and  the  like,  should  be  divided  or  thinned  down 
with  a  knife,  but  in  such  a  manner  that  the  original  shape  can 
be  easily  made  out. 

785.  Carices  should  be  always  collected  when  the  fruit  is  full- 
grown,  but  not  so  ripe  as  to  fall  away.     So  also  should  other 
Cyperaceae ;  }-et  it  is  well  to  collect  also  earlier  specimens  of 
these  in  flower.     Grasses,  on  the  other  hand,  should  generally 
be  collected  soon  after  they  come   into   blossom.      For  when 
Imature  the  spikelets  in  many  species  break  up  and  fall  away  in 
drying.      The  culm,  leaves,  and  root  of  Sedges  and  Grasses 
should  be  preserved,  as  well  as  the  inflorescence.     The  root  is 
no  less  important.     Cespitose  species  should  be  so  collected  and 
preserved  as  to  show  the  tufted  character.     The  culms  of  most 
sedges  and  grasses  act  stubbornly  when  bent  for  arrangement 
in  portfolio  or  press,  and  are  not  disposed  to  stay  in  place.    This 
difficulty  is  promptly  remedied  by  crushing  with  the  teeth  the 
angles  made  by  the  bending.     Or  these  may  be  thrust  through 
slits  of  paper.     In  drj'ing  Sedges  and  Grasses,  very  moderate 
pressure  should  be  employed. 


HERBORIZATION.  375 

786.  Some  aquatic  plants  (Algae  especially)  are  so  soft  and 
flaccid  that,  to  secure  them  in  their  proper  shape,  they  must  be 
placed  in  clear  water  and  floated  out  by  inserting  beneath  them 
the  paper  on  which  they  are  to  remain  permanently,  either  the 
regular  mounting  paper,  or  a  thinner  white  paper  which  when 
dry  can  be  pasted  on  the  herbarium  sheet.     If  likely  to  adhere 
to  the  sheet  or  drier  above  them  in  the  press,  a  piece  of  oiled  or 
stearine  paper  may  be  laid  directly  on  the  specimens  to  prevent 
their  sticking.     Also  viscous  or  glutinous  plants  which  are  liable 
to  adhere  to  the  sheets  enclosing  them  may  be  sprinkled  with 
Lycopodium  spores,  powdered  soapstone,  or  some  similar  sub- 
stance. 

787.  The  name  of  the  plant  if  known,  but  by  all  means  the 
locality  and  date  of  collection,  with  any  other  descriptive  re- 
marks regarded  necessary,  should  be  written  on  a  ticket  or  on 
the  sheet  when  it  is  put  into  the  press.     Never  omit  to  record 
the  time  and  place  of  collection,  as  a  specimen  of  unknown  date 
and  locality  loses  much  of  its  value  and  interest. 

788.  Drying  Specimens.     The  chief  requisite  for  good  herba- 
rium specimens  is  the  extraction  of  the  moisture  from  the  green 
plant   as  rapidly  as  possible  under  a  pressure  which  obviates 
brittleness.     This   is  to  be  affected  by  placing  the  thin  sheets 
containing  the  specimens  between  layers  of  bibulous  paper,  called 
driers,  and   applying   moderately  strong  pressure  to  the   pile. 
For  driers   nothing   can   be  better  than  thick  blotting  paper, 
except  that  it  is  too  expensive,  and  the  same  may  be  said  of 
an  English  drying  paper  made  for  the  purpose.    Equally  good 
driers  are  made  of  the  thick  and  felt-like  brown  paper  which, 
after  saturation  with   coal-tar,  is  here  largely  used  under  the 
clapboards  of  wooden  houses  and  under  slate-roofing.     It  is  a 
cheap  material,  and  is  to  be  obtained,  cut  into  sheets  of  18  by 
12  inches.     Or  driers  may  be  made  of  old  newspapers  or  of  any 
soft  wrapping   paper,  cut   or   folded   to   the   proper   size,  and 
stitched  (very  expeditiously  by  a  sewing-machine),  or  joined  by 
e3-elet  paper-fasteners  at  two  corners,  in  packages  of  a  dozen  or 
more  leaves  to  a  drier.    It  is  well  to  have  a  large  supply  of  driers 
and  specimen-sheets  ready  for  use. 

789.  A  half  dozen  or  more  pieces  of  thin  boards,  18  inches 
long  and  12  inches  wide,  should  be  provided.     They  are  used 
at  the  top  and  the  bottom  of  the  pile  when  pressing,  and  also 
for  dividing  it  into  suitable  sections,  especially  for  separating 
the  packages  of  plants  which  were  put  into  press  at  different 
periods,  and  dividing  up  these  packages  themselves,  if  too  large. 
For  the  plants  dry  better  in  small  sections  and  with  the  pressure 


376  PHYTOGRAPHY. 

evenly  distributed.  Hence  it  is  best  to  have  these  sections  not 
over  five  or  six  inches  in  thickness,  nor  should  the  pile  itself 
be  carried  too  high,  never  exceeding  two  feet.  Painted  binders' 
boards  may  be  used,  instead  of  the  common  boards,  to  separate 
interior  divisions.  Some  botanists  use  a  kind  of  lattice  made  of 
two  layers  of  thin  strips  or  laths,  crossing  each  other.  This  is 
said  to  allow  free  escape  of  the  moisture  by  evaporation,  and  so 
to  accelerate  drying,  as  in  the  case  of  the  wire  press. 

790.  For  giving  pressure,  various  ways  have  been  contrived. 
The  Screw-press  is  convenient  and  compact,  but  objectionable, 
because  it  does  not  follow  up  the  pressure  as  the  plants  shrink 
in  drying.    This  objection  does  not  apply  to  the  Lever-presses,  but 
they  are  usually  unwieldy.     Fortunately,  one  of  the  best  forms 
of  the  drying-press,  as  well  as  the  simplest  and   cheapest,  is 
merely  a  board  with  weights  placed  on  the  top  of  the  pile  of 
specimens.     Here  the  pressure  is  continuous,  constantly  follow- 
ing the  shrinkage  of  the  plants.     The  weight  on  a  pile  should 
vary  from  25  to  100   pounds,  according  to  the  nature  of  the 
specimens  and  the  quantity  in  the  press.     On  an  average,  60 
pounds  is  sufficient  for  most  plants.     If  much  greater  pressure 
is  used,  there  is  danger  of  crushing  the  more  delicate  parts  of  the 
specimen,  and  thereby  impairing  its  scientific  value.    For  weights, 
bars  or  masses  of  iron  may  be  used,  boxes  filled  with  sand, 
stones,  and  the  like.  • 

791.  Specimens  brought  home  in  the  botanical  box  must  be 
placed  in  such  thin  specimen-sheets  as  are  used  in  portfolio. 
In  putting  plants   in  specimen- sheets,  whether  in  portfolio  or 
press,  it  is  well  to  take  some  pains  to  spread  out  the  specimens 
neatly  ;  for  a  little  care  now  may  save  much  later  labor.     How- 
ever, with  most  species,   any  carelessness  in  this  respect  can 
be  remedied  at  the  first  change  of  driers.    But  there  are  some 
plants,  previously  referred  to,  so  peculiarly  sensitive  that  what- 
ever adjustment  they  receive  must  be  given  at  the  time  they  are 
first  placed  in  their  sheets. 

792.  Although  plants  can,  if  necessary,  be  kept  fresh  for  several 
days  in  box  or  portfolio,  on  returning  from  a  collecting  trip  they 
should  be  transferred  to  the  home  press  as  early  as  possible.     In 
the  transference,  particular  care  should  be  taken  to  straighten 
out  and  remove  all  folds  and  crumpling  of  the  leaves,  petals, 
fronds,  &c.,  and  to  arrange  the  specimen  as  naturally  as  possi- 
ble, so  as  to  show  the  proper  habit.     Both  sides  of  the  flowers 
and  leaves  should  be  exhibited.     Plants  that  were  put  directly 
into   press   should   receive   this   special   attention   at   the   first 
change  of  driers,  which  on  this  account  should  be  made  within 


HEEBORIZATION.  377 

several  hours  afterwards.  The  stubbornness  and  elasticity,  so 
troublesome  in  specimens  when  first  put  in,  will  then  have 
mostly  disappeared,  and  the  whole  specimen  will  be  found  suffi- 
ciently flaccid  to  have  every  part  stay  as  arranged.  If  this  first 
change  is  deferred  longer  than  ten  or  twelve  hours,  the  speci- 
mens of  many  species  become  too  dry  for  making  the  alterations 
required.  At  this  time  small  pieces  of  bibulous  paper  may  be 
placed  between  leaves,  or  other  portions  of  the  plant  which  over- 
lap, to  prevent  moulding  or  discoloration,  and  to  hasten  dr}ang. 
It  is  well  to  change  these  fragments  of  paper  with  the  driers  for  the 
first  day  or  two  :  afterwards  they  may  remain  with  no  detriment. 

793.  To  have  the  specimens  retain  their  natural  color  and 
general  appearance,  they  should  be  dried  as  rapidly  as  possible ; 
and  this  result  is  best  secured  by  frequent  changing  of  the  driers. 
These  should  be  changed  at  least  once  a  day  for  the  first  four  or 
five  days,  and  afterwards  every  other  da}-,  until  the  specimens 
are  thoroughly  dried.     But  a  marked  improvement  in  the  speci- 
mens will  result  from  more  frequent  changes   during   the  first 
day  or  two.    The  first  day  with  Grasses,  Sedges,  and  their  allies, 
and  the  first  two  days  with  most  other  plants,  are  of  more  impor- 
tance than  all  the  subsequent  time.    As  an  experienced  collector 
declares  :  "  Two  or  three  changes  of  the  driers  during  the  first 
twenty-four  hours  will  accomplish  more  than  a  dozen  changes 
after  the  lapse  of  several  days.     The  most  perfect  preservation 
of  the  beautiful  colors  of  some  Orchids  has  been  effected  by 
heating  the  driers  and  changing  them  every  two  hours  during 
the  first  day." 

794.  Heated  driers  are  very  efficient ;  and  the  best  mode  of 
heating  is  to  expose  them  to  the  sunshine,  and  bringing  them  in 
hot  to  make  the  change  at  once,  or  as  soon  as  possible. 

795.  The  number  of  driers  interposed  between  the  specimen- 
sheets  should  depend  upon  the  plants  and  the  frequency  of  the 
changes  :  two  will  suffice  when  the  driers  are  changed  very  often  ; 
but  more  must  be  employed  when  the  plants  are  thick  and  succu- 
lent.    Uniform  pressure  may  be  secured  with  large  and  coarse 
plants  by  placing  strips  of  pasteboard  or  pieces  of  cotton-bat- 
ting about  the  sides  of  the  package.     Ringlets  of  cotton  may  be 
placed  about  some  of  the  larger  flower-heads  of  the  Compositse,  &c. 

796.  The  time  required  to  dry  specimens  varies  with  different 
species  and  with  the  season :  it  depends  also  on  the  frequency 
of  the  changes  and  the  temperature  of  the  driers.     By  changing 
daily,  the  time  is  usually  from  four  or  five  days  to  a  week.    But, 
with  two  changes  a  day  for  the  first  day  or  two  and  with  heated 
driers,  the  process  may  be  completed  in  half  the  usual  time,  ana 


378  PHYTOGRAPHY. 

the  specimens  will  be  in  much  finer  condition.  An  experienced 
collector  has  no  difficult}7  in  ascertaining  whether  a  plant  is  com- 
pletely cured  or  not,  while  to  a  novice  it  is  often  a  matter  of 
uncertainty.  A  thoroughly  dried  plant  can  be  usually  told  by 
its  peculiar  hay-like  rattle  when  disturbed  ;  also  by  placing  the 
plant  against  the  cheek.  If  there  is  a  sensation  of  coolness,  the 
plant  is  still  moist. 

797.  If  the  thick  leaves  of  fleshy  plants  are  immersed  for 
a  few  moments  in  hot  water,  the  period  of  desiccation  will  be 
greatly  hastened ;    but  they  frequently  turn   dark  as   a  conse- 
quence  of  the  immersion.      The   drying   of  such   plants,  and 
particularly  of  the  Monocotyledons,  may  be  advantageously  ex- 
pedited by  placing  them  between  several  driers  and  ironing  them 
with  hot  irons.     Small  plants  may  be  very  neatly  dried  in  old 
books.     Very  beautiful  specimens  may  be  made  by  placing  the 
plant  in  a  tall  and  narrow  vessel,  and  pouring  over  it  a  sufficient 
quantity  of  clean  and  dry  sand.    When  the  moisture  is  absorbed, 
it  may  be  flattened  in  a  press. 

798.  In  shifting  the  driers  of  a  collection,  place  the  package 
to  be  changed  at  the  left  hand  on  the  table  or  counter,  the  new 
pile  in  front  with  its  length  parallel  to  the  person,  —  a  position 
the  most  favorable  for  giving  any  needed  attention  in  arranging 
specimens,  —  while  fresh  driers  may  be  placed  at  the  right  hand, 
or  beyond  the  pile  in  front.    Thus  arranged,  the  sheets  of  speci- 
mens can  be  rapidly  shifted  into  their  fresh  driers. 

799.  The  moist  driers  may  be  spread  out  in  the  sunshine  to 
dry,  or  strung  on  a  line  in  a  warm  room,  or  in  the  open  air,  if 
not  too  windy.     Very  moist  driers   may  be   thorough^  dried 
within  an  hour,  if  spread  in  the  hot  sunshine.     In  inclement 
weather,  they  must  be  dried  by  the  fire. 

800.  To  recapitulate  the  most  important  points  in  good  speci- 
men-making :    Use   specimen-sheets  to   hold  the  plants  undis- 
turbed during  the  whole  process  of  drying :  use  plenty  of  the 
most  bibulous  driers,  sun-dried  and  heated  when  practicable  :  do 
not  make  the  piles  too  large  :  make  the  first  shift  of  driers  within 
a  few  hours,  at  that  time  making  all  needed  adjustment  of  the 
flaccid  specimens :  change  the  driers  twice  a  day  for  the  first 
day  or  two. 

801.  For  collecting  and  preserving  specimens  on  a  journey, 
or  when  moving  from  place  to  place,  some  modification  of  the 
stationary  press  is  requisite.    The  Travelling-press  must  be  porta- 
ble :  accordingly  the  pressure  is  applied  by  strong  leather  straps 
with  buckles.     There  should  be  three  straps,  one  girding  the 
package  around  each  end,  and  one  lengthwise.     The  top  and 


HERBORIZATION.  379 

bottom,  if  of  thin  boards,  must  be  cleated.  or  compounded  of 
wood  with  the  grain  in  opposite  directions,  or  very  stout  binder's 
board  or  trunk-board  may.  be  advantageously  used.  This  should 
be  covered  with  coarse  cotton  or  linen  cloth,  glued  fast  and  well 
painted.  While  stationary,  the  pressure  may  be  giveorfoy  means 
of  weights  when  more  convenient. 

802.  The  Wire  press,  now  much  in  use,  is  a  press  of  this  porta- 
ble kind,  in  which  the  boards  are  replaced  by  sheets  of  wire  net- 
ting, with  wide  meshes,  and  surrounded  and  strengthened  by  a 
strong  but  light  iron  border.     Straps  with  buckles  are  used  to 
hold  the  parts  and  contents  together  and  to  apply  the  pressure, 
as  in  the  ordinary  travelling-press.     Besides  its  portability,  the 
advantages  of  such  wire  presses  are  that,  in  a  small  way,  they 
may  serve  both  as  portfolio  for  collecting  and  as  press  for  dry- 
ing ;  also  that,  as  the  drying  takes  place  mainly  by  evaporation 
instead  of  absorption,  much  less  paper  is  required,  and  the  trouble 
of  changing  the  driers  is  saved. l    In  fair  weather,  the  press  filled 
with  plants  may  be  hung  in  the  wind  or  sunshine,  in  foul  weather 
near  a  fire.     The  disadvantage  is  that  specimens  dried  in  this 
way  are  apt  to  be  brittle.     To  use  this  sj'stem  advantageously, 
the  botanist  should  have  at  least  two  such  presses  in  operation, 
one  for  collecting,  while  the  other  is  in  use  for  drying. 

803.  Poisoning.     Dried  specimens  are  liable  to  the  depreda- 
tions of  certain  insects,  especially  of  their  larvae.     The  principal 
pest  is  a  small  brown  beetle,  Anobium  paniceum,  L.  ;   the  perfect 
insect  does  considerable  damage,  the  larva  vastly  more.     Plants 
with  milky  juice,  such  as  Asclepiadese,  Apocynaceae,  and  Eu- 
phorbiaceae,  those  containing  bitter  principles,  such  as  Gentians 
and  Willows,  and  generally  such  plants  or  such  organs  as  con- 
tain much  protoplasm  or  azotized  matter,  are  most  subject  to 
attack.    Ranunculaceae,  Umbelliferae,  and  Compositae  are  seldom 
spared  ;  while  Labiatae  mostly  escape,  probably  on  account  of  the 
volatile  oil  which  they  contain.    Even  Ferns  are  liable  to  have  the 
parts  of  fructification  eaten  away.     To  a  certain  extent,  the  im- 
pregnation of  the  herbarium-cases  with  camphor,  naphthaline, 
or  strong-scented  oils,  may  exclude  the  vermin.     But  safety  is 
secured  only  by  poisoning. 

804.  The  proper  poison  is  corrosive  sublimate,  dissolved  in 
strong  (95  per  cent.)  alcohol.    Drop  into  the  alcohol  as  much  cor- 

1  Prof.  A.  Wood  seems  to  have  been  the  first  to  call  the  attention  of 
American  botanists  to  this  system,  which  he  has  earnestly  advocated. 

An  improved  form  of  this  wire  press,  well  adapted  both  for  collecting 
and  pressing  in  moderate  quantity,  is  made  and  sold,  at  a  small  price,  by 
Paul  Rcessler,  optician,  at  New  Haven,  Connecticut. 


380  PHYTOGEAPHY. 

rosive  sublimate  as  it  will  take  up,  then  add  a  trifle  more  of  alcohol, 
so  as  to  keep  the  solution  just  below  the  point  of  saturation. 
The  stronger  the  solution  the  better,  except  that,  at  full  satura- 
tion and  where  copiously  used,  an  efflorescence  may  sometimes  be 
left  on  the  surface  of  the  poisoned  specimens  upon  the  evapora- 
tion of  the  alcohol.  Some  add  to  the  solution  some  carbolic 
acid,  at  the  rate  of  a  fluid  ounce  to  each  quart  of  alcohol.  The 
solution  may  be  applied  with  a  soft  brush  (one  with  no  metal  in 
its  fastening) ,  or  by  a  dropping  bottle,  or  even  the  specimens 
may  be  dipped  in  the  solution  placed  in  a  flat  porcelain  dish. 
The  brush  (using  a  pretty  large  and  soft  one)  is  the  most  con- 
venient and  efficient.  The  moistened  specimens  should  be  placed 
between  driers  and  in  shallow  piles  until  the  alcohol  evaporates. 
.  805.  Thoroughly  poison  all  specimens  before  admitting  them 
to  the  herbarium.  It  is  well  to  poison  all  specimens  whatever, 
as  soon  as  they  are  made  or  at  the  close  of  the  botanizing  sea- 
son, as  well  those  intended  for  exchanges  as  for  the  collector's 
own  herbarium. 

806.  Keep  all  specimens  between  sheets  of  paper,  or  within 
folded  sheets,  not  too  crowded  or  overlaid,  away  from  dust,  and 
in  a  perfectly  dry  place,  so  as  to  avoid  mould.     When  attacked 
by  mould,  the  corrosive- sublimate  solution  should  be  applied. 
A  properly  dried  specimen,  duly  cared  for,  should  be  as  lasting 
as  the  paper  which  holds  it. 

807.  The   Herbarium,  called  by  the   earlier  botanists  Hortus 
Siccus,  is  a  collection  of  dried  specimens,  named  and  systemat- 
ically arranged.     It  is  indispensable  to  the  working  systematic 
botanist,  and  every  devotee  of  botany  should  possess,  or  have 
access  to  an  herbarium  containing  representatives  of  the  plants 
of  the  immediate  vicinity  or  district,  if  not  of  the  whole  country. 
Or  an   herbarium  may  be  restricted  to  a  particular  family  of 
plants,  made  the  object  of  special  study.     A  general  herbarium 
should  contain  specimens  representing  all  the  natural  orders  and 
as  many  of  their  genera  and  species  as  possible. 

808.  The  form  of  the  herbarium  as  to  the  size  of  its  sheets  is 
considerably  variable.    That  of  Linnaeus  is  of  the  size  of  foolscap 
paper:    this  would   now  be  universally  regarded  as  much  too 
small.     The  principal  British  herbaria  adopt  the  size  of  16£  by 
10^  inches,  which  is  rather  too  narrow,  rarely  permitting  two 
specimens  of  the  same  species  of  any  considerable  size  to  be 
placed  side  by  side  on  the  same  sheet.     In  the  United  States, 
16£  inches  in  length  by  1  If  in  width  is  adopted  ;  that  is,  for  the 
genus-covers,  the  species-paper  being  a  quarter  of  an  inch  nar- 


THE  HERBARIUM.  381 

809.  The  specimens  representing  each  species  may  either  be 
laid  within  a  doubled  sheet,  loosely  (as  in  some  European  her- 
baria), or  fastened  in  place  by  narrow  slips  of  gummed  paper 
(which  is  much  better) ,  or  else  they  may  be  glued  bodily  to 
single  sheets  of  strong  and  stiff  white  paper. 

810.  The  former  is  an  excellent  plan  for  a  limited  collection. 
It  is  an  advantage  that  a  specimen  can  be  taken  up  and  examined 
on  all  sides  ;  also,  that  indifferent  specimens  can  at  any  time  be 
exchanged  for  better  ones.     But  a  large  herbarium  on  this  plan 
becomes  cumbrous   and   inconvenient  for  ready  reference  and 
comparison. 

811.  The  best  plan  in  a  large  herbarium,  and  one  much  to  be 
consulted,  is  to  attach  the  specimens  completely,  by  any  kind 
of  strong  and  light-colored  glue,  to  single  sheets,  or  rather  half 
sheets.     The  specimens  are  thus  safe  from  injury  under  reason- 
able handling,  and  can  be  turned  over  and  examined  with  as 
much  facility  as  a  series  of  maps  or  engravings.     The  species- 
paper  should  be  of  writing-paper  stock,  or  of  equal  firmness,  of 
compact  texture,  well  sized  and  calendered,  and  of  a  weight  in 
size  of  16£  by  11 J  inches  of  about  18  pounds  to  the  ream  of 
480  flat  sheets.     The  paper  should  be  furnished  square-cut  on 
all  sides,  in  the  manner  of  "flat  cap."     Stiffness  is  the  great 
desideratum. 

812.  In  no  case  should  more  than  one  species  be  knowingly 
attached  to  the  same  sheet.    But  of  very  many  species  there  will 
be  room  for  more  than  one  specimen.     And  specimens  from  dif- 
ferent localities,  of  different  forms,  and  in  various  stages  of 
flowering  and  fructification,  are  always  desirable.    The  full  name 
of  the  plant  should  be  written  at  the  lower  right-hand  corner  of 
the  sheet,  or  a  ticket  should  there  be  attached  by  glue  or  traga- 
canth  paste.     Each  specimen  should  have  its   ticket,  similarly 
attached,  or  a  memorandum  upon  the  sheet,  indicating  the  hab- 
itat or  the  special  locality,  date  of  collection,  name  of  collector, 
and  any  other  desirable  information  which  the  specimens  them- 
selves do  not  furnish.    When  there  are  loose  flowers  or  fruits,  or 
when  any  of  these  have  been  detached  for  dissection  and  micro- 
scopical investigation,  it  is  well  to  preserve  them,  placing  them 
in  little  paper  pockets  or  envelopes  and  pasting  these  upon  the 
sheet  close  to  the  specimen  to  which  they  pertain.     Sketches  of 
parts  dissected  may  be  drawn  upon  the  sheet.     Notes  and  mem- 
oranda received  with  the  specimen  or  too  extended  to  be  entered 
upon  the  sheet  may  be  folded,  inserted  in  such  envelopes,  and 
made  fast  to  the  sheet.     Many  botanical  collections  are  distrib- 
uted with  printed  tickets.     These,  and  all  authenticating  tickets 


382  PHYTOGRAPHY. 

or  notes,  should  be  attached  to  the  sheet  near  to  the  specimen 
they  belong  to.  In  view  of  this,  printed  and  written  tickets 
should  be  of  small  size.1  A  ticket  which  exceeds  four  by  two 

1  All  printing  on  an  herbarium  ticket  should  be  in  plain  type ;  and  fancy 
borders,  uselessly  occupying  room,  should  be  avoided.  If  any  border  is 
thought  needful,  it  should  be  of  plain  lines.  It  is  not  desirable  to  parcel  out 
the  space  on  a  ticket  with  separate  lines  and  headings  for  habitat,  date  of 
collection,  time  of  flowering  or  fruiting,  name  of  collector,  and  the  like. 
These  particulars  may  conveniently  be  entered  at  the  bottom  or  top  of  the 
ticket,  as  may  be  convenient,  leaving  the  rest  of  the  space  free  for  the  name 
of  the  plant,  the  authority,  and  perhaps  a  synonym. 

Tickets  for  specimens  distributed  among  other  botanists  may  well  have  a 
head-line  indicating  the  source,  such  as  "  EX  COLL.  c.  c.  PARRY,"  or  "  EX  HERB." 
or,  in  English,  "  FROM  THE  HERBARIUM  OF  "  the  botanist  who  communicates 
the  specimen.  The  following  may  serve  as  an  example  of  a  simple  ticket 
for  the  sending  out  of  dried  specimens,  and  of  the  way  in  which  the  ticket 
may  be  filled  out  with  the  name  of  the  plant,  its  habitat  and  station,  name 
of  collector  and  time  of  collection. 


EX  HERB.  A.  GRAY. 


For  the  botanist's  own  herbarium,  it  is  well  to  use  a  blank  ticket  with  a 
printed  heading  like  the  specimen  above,  but  with  the  "  ex  "  omitted. 

When  a  considerable  collection  is  made  in  any  particular  botanical  explo- 
ration or  excursion,  and  numerous  or  several  specimens  of  the  same  species 
are  gathered,  to  be  distributed  among  botanists  in  the  way  of  exchange  or 
otherwise,  these  are  commonly  given  out  under  numbers  and  with  a  printed 
heading  to  a  special  ticket.  The  following  is  an  approved  form  of  such  a 
ticket,  and  of  the  mode  in  which  it  may  be  filled  up  in  writing  by  inserting 
the  name  of  the  species,  the  locality,  &c. 


ALPINE  FLORA  OF  THE  ROCKY  MOUNTAINS. 

Coll.  C.  C.  PARRY.    1872. 

No. 


•Wrvf/w  ,  XTCuO  , 


COLORADO, 

<HX.  <«-*  > 


THE   HERBARIUM.  383 

inches  is  a  nuisance ;  and  those  of  an  inch  and  a  quarter  or  an 
inch  and  a  half  in  width  and  three  or  four  inches  in  length  are 
most  commodious. 

813.  The  sheets  of  all  the  species  of  the  same  genus,  when 
not  too  numerous,  or  of  a  particular  section  of  it,  or  any  conven- 
ient number,  should  be  consigned  to  one  genus-cover.     The  best 
genus-covers  are  of  manilla-rope  paper,  the  "  bleached  manilla  " 
such  as  that  of  which  tags  are  made  is  the  neatest  article,  but 
rather  more  expensive  :  they  are  in  whole  or  folded  sheets  (pref- 
erably in  quarter  quires),  accurately  trimmed  at  top,  bottom, 
and  front  edge  to  the  size  of  16 \  by  11|  inches;  that  is,  the 
folded  sheet  as  used  is  a  trifle  longer  and  a  quarter  of  an  inch 
wider  than  the  species-sheets  it  holds.     The  sheets  to  be  firm 
enough  should  weigh  1|  or  1|  ounces  each,  or  from  45  to  52 
pounds  the  ream.    The  generic  name  should  be  written  in  a  bold 
hand  on  the  lower  left-hand  corner ;  that  is,  on  the  upper  face  next 
the  back :  at  or  near  the  lower  right-hand  corner,  the  name  of  the 
contained  species  may  be  written  either  with  a  pencil  or  in  ink. 

814.  The  genera  should  be  arranged  in  the  herbarium  accord- 
ing to  some  systematic  work,  and  numbered  accordingly  on  the 
covers. 

815.  The  herbarium  must  be  preserved  in  close  cabinets  or 
cases  free  from  the  access  of  dust.     Tin  cases,  just  deep  and 
wide  enough  to  receive  comfortably  the  genus-covers,  and  about 
six  inches  high,  the  hinged  lid  being  one  end,  may  be  recom- 
mended  for  a  small   collection,  as  they  are   dust  and  insect 
proof,  are  portable,  and  may  readily  be  arranged  on  shelves. 
But,  for  any  herbarium  of  considerable  size  and  continued  growth, 
wooden  cabinets  with  well-fitted  doors  are  to  be  preferred  ;  the 
interior  of  the  cabinets  being  divided  into  pigeon-hole  compart- 
ments, fully  12  inches  wide  in  the  clear  and  17  inches  deep,  and 
not  over  6  inches  or  in  small  herbaria  not  over  4  inches  high. 
Into  such  pigeon-holes,  the  genus-covers  with  their  contents  will 
slide  readily,  and  ma}'  be  compactly  stowed  away.     An  index 
to  the  genera  of  each  order  may  be  affixed  to  the  interior  of  the 
cabinet  doors,  or  pasted  upon  the  upper  face  of  thin  boards, 
inserted  at  the  beginning  of  each  order.     The  name  of  the  order, 
written  or  printed  in  bold  letters,  may  be  pasted  upon  the  front 
edge  of  this  board,  or  upon  a  flap  of  card-board  affixed  to  it. 
Moreover,  it  is  well  to  write  the  name  of  the  order  upon  each 
genus-cover. 

816.  Except  in  public  collections,  where  fixed  cases  may  be' 
preferred,  the  cabinets  should  individually  be  small,  only  three 
or  four  feet  high,  and  containing  only  two  or  four  vertical  rows 


PHYTOGKAPHY. 


of  compartments.  Such  cabinets  can  be  increased  in  number  as 
required,  are  portable,  and  can  be  disposed  in  any  order,  side  by 
side  or  one  surmounting  another,  as  may  be  most  convenient. 
The  doors  should  be  so  constructed  as  to  open  and  shut  readily, 
but  to  close  tightly,  so  as  to  exclude  dust  and  insects.1 


1  An  excellent  plan  for  small  and  inexpensive  herbarium  cabinets,  of  a 
portable  character,  is  proposed  and  illustrated  by  Dr.  Parry,  in  the  American 
Naturalist,  viii.  471.  Each  small  case  is  in  fact  a  plain  wooden  box,  wide 
enough  to  hold  two  tiers  of  pigeon-hole  compartments,  and  of  any  desirable 
height  (three  compartments  high  in  Dr.  Parry's  plan,  but  double  the  number 
might  be  better)  :  the  entire  front  consists  of  a  pair  of  doors  meeting  in  the 
centre,  there  fastened  by  a  flush  spring  catch ;  the  doors  bevelled  on  the  inside, 
with  a  corresponding  bevel  on  the  case,  to  which  they  are  attached  by  out- 
side hinges,  so  that  in  opening  at  a  right  angle  there  are  no  sharp  corners  to 
hinder  the  drawing  out  of  the  herbarium  papers ;  also  allowing  the  cases 
to  stand  close  side  by  side,  as  well  as  one  upon  another,  without  interfering 
with  the  free  opening  of  the  doors.  These,  moreover,  may  swing  quite  back 
against  the  sides  without  in  any  way  straining  the  hinges.  For  lifting,  a 
pair  of  flush  handles,  countersunk  to  the  level  of  the  wood,  may  be  attached 
to  the  sides.  When  the  herbarium  has  to  be  removed  to  a  distant  place, 
these  cases,  having  no  projecting  knobs  or  handles,  will  go  readily  into  ordi- 
nary packing-boxes. 


ABBREVIATIONS. 


1.  OF  NAMES  OF  BOTANISTS  AND  BOTANICAL  AUTHORS. 

Ach.    = 

Acharius. 

Berth.  = 

Berkhey. 

Adans. 

Adanson. 

Berken. 

Berkenhout. 

Afz. 

Afzelius. 

Berland. 

Berlandier. 

Ag. 

Agardh. 

Bernh. 

Bernhardi. 

C.Ag. 

C.  A.  Agardh. 

Bert. 

Bertero. 

J.Ag. 
Ag.f. 

>  J.  G.  Agardh,  son. 

Bertol. 
Bess. 

Bertoloni. 
Besser. 

Ait. 

Alton. 

Bieb. 

Marschall  von  Bieberstein. 

All. 

Allioni. 

Bigd. 

Jacob  Bigelow. 

Amm. 

Amman. 

Bisch. 

Bischoff. 

Anders. 

Andersson  of  Stockholm. 

Bahm. 

Boehmer. 

Andr. 

Andrews. 

Boerh. 

Boerhaave. 

Andrz. 

Andrzejowski. 

Boiss. 

Boissier. 

Aresch. 

Areschoug. 

Boland. 

Bolander. 

Am. 

Arnott. 

Bong. 

Bongard. 

Arrh. 

Arrhenius. 

Bonpl. 

Bonpland. 

Asch. 

Ascherson. 

Bork. 

Borkhausen. 

Aubl. 

Aublet. 

Borsz. 

Borszcow. 

Brack. 

Wm.  D.  Brackenridge. 

Bab. 

Babington. 

Brebis. 

Brebisson. 

Bail. 

Baillon. 

Bref. 

Brefeld. 

Ball. 

Balbis. 

Brew.S?   \ 

W.  H.  Brewer  &  Sereno 

Baldw. 

Baldwin. 

Wats.    1 

Watson. 

Balf. 

Balfour. 

Brid. 

Bridel. 

Barn. 

Barneoud. 

Brong. 

Brongniart. 

Barr. 

Barrelier. 

Brot. 

Brotero. 

Bart. 

Benj.  Smith  Barton. 

Brouss. 

Broussonet. 

W  Bart. 

Wm-  P.  C.  Barton,  nephew. 

Br.,  R.  Br.  Robert  Brown. 

Bartr. 

John  Bartram. 

P.  Br. 

Patrick  Browne. 

Bartr.f. 

Wm.  Bartram. 

Brunf. 

Brunfels. 

Bauh. 

Bauhin. 

Buckl. 

Buckley. 

Beauv. 

Palisot  de  Beauvois. 

Bull. 

Bulliard. 

Benj. 

Benjamin. 

Burm. 

Burman. 

Benn. 

J  J.  Bennett. 

Buxb. 

Buxbaum. 

A.  Benn. 

A.  W.  Bennett. 

Benth. 

Bentham. 

Cam. 

Camerarius. 

Berg. 
Berk. 

Bergius. 
M.  J.  Berkeley, 

Camb.       \ 
Cambes.    \ 

Cambessedes. 

386 


ABBREVIATIONS. 


Campd.  = 

Campdera. 

Eat.    = 

Amos  Eaton. 

Cand. 

DeCandolle,  usually  DC. 

D.  C.  Eat. 

D.  C.  Eaton,  grandson. 

Casp. 

Caspary. 

Edgew. 

Edgeworth. 

Cass. 

Cassini. 

Edw. 

Edwards. 

Catesb. 

Catesby. 

Ehren. 

Ehrenberg. 

Cav. 

Cavanilles. 

Ehrh. 

Ehrhart. 

Cerv. 

Cervantes. 

Eichl. 

Eichler. 

Cham. 

Chamisso. 

Eiseng. 

Eisengrein. 

Chapm. 

A.  W.  Chapman. 

Eli. 

Elliott. 

Chao. 

Chavannes. 

Endl. 

Endlicher. 

Chois. 

Choisy. 

Engelm. 

Engelmann. 

Clayt. 

Clayton. 

Engl. 

Engler. 

Clus. 

Clusius. 

Eschs. 

Eschscholtz. 

Collad. 

CoUadon. 

Eschw. 

Eschweiler. 

Cdm. 

Colmeiro. 

Ettingsh. 

Ettingshausen. 

Comm. 

Commelin. 

Corn. 

Cornuti. 

Fendl. 

Fendler. 

Coss. 

Cosson. 

Feuil. 

Feuille'e. 

Cunn. 

Cunningham,  A.  or  J. 

Fingerh. 

Fingerh  uth. 

Curt. 

Wm.  Curtis. 

Fisch. 

Fischer. 

M.A.Curt.  M.  A.  Curtis. 

Forsk. 

Forskll. 

Forst. 

Forster. 

Dalech. 

Dalechamps. 

Fourn. 

Fournier. 

Dalib. 

Dalibard. 

Fresen. 

Fresenius. 

Darl. 

Darlington. 

Freyc. 

Freycinet. 

DC.          ) 

Fred. 

Frcelich. 

1 

DeC.        j 

A.  P.  DeCandolle. 

A.  DC. 

Alphonse  DeCandolle,son. 

GcErtn. 

J.  Gaertner. 

Cos.  DC. 

Casimir  DeCandolle,  the 

GcRrtn.f. 

C.  T.  Gaertner. 

Decne. 

Decaisne.          [grandson. 

Gardn. 

Gardner. 

Deless. 

Delessert. 

Garid. 

Garidel. 

Dennst. 

Dennstedt. 

Gasp. 

Gasparrini. 

Desc. 

Descourtilz. 

Gaud. 

Gaudin. 

Desf. 

Desfontaines. 

Gaudich. 

Gaudichaud. 

Desj. 

Desjardins. 

Germ. 

Germain. 

Desmar. 

Desmazieres. 

Gesn. 

Gesner. 

Desmoul. 

Desmoulins. 

Gilib. 

Gilibert. 

Desv. 

Desvaux. 

Ging. 

Gingins  de  Lassaraz. 

Dicks. 

Dickson. 

Gis. 

Giseke. 

Diesb. 

Diesbach. 

Gled. 

Gleditsch. 

Dieter. 

Dieterich. 

Gleich. 

Gleichen. 

Dietr. 

Dietrich. 

Glox. 

Gloxin. 

Dill. 

Dillenius. 

Gmel. 

J.  G.  Gmelin. 

Dittw. 

Dillwyn. 

C.  Gmel. 

C.  C.  Gmelin  of  Baden. 

Dod. 

Dodonaeus  (Dodoens). 

S.  Gmel. 

S.  G.  Gmelin. 

D'Orb. 

D'Orbigny. 

Godr. 

Godron. 

Dorst. 

Dorstenius. 

Gcepp. 

Goeppert. 

Dougl. 

Douglas. 

Good. 

Goodenough. 

Drej. 

Drejer. 

Gren. 

Grenier. 

Dryand. 

Dryander. 

Grev. 

Greville. 

Dufr. 

Dufresne. 

Griseb. 

Grisebach. 

Duham. 

Duhamel  du  Monceau. 

Gram. 

Greenland. 

Dumort. 

Dumortier. 

Gran.        \ 

Gronovius. 

Dun. 

Dunal. 

Gronov.    ! 

ABBREVIATIONS. 


38T 


Guett.    =  Guettard. 

Jacq.     = 

N.  J.  Jacquin. 

Guib.          Guibord. 

Jacq.f. 

J.  F.  Jacquin,  son. 

Guiliem.      Guillemin. 

J.  St.  Hil 

Jaume  St.  ililaire. 

Guimp.       Guimpel. 

Jord. 

Jordan. 

Gunn.         Gunnerus. 

Jungh. 

Junghuhn. 

Guss.          Gussone. 

Juss. 

A.  L.  Jussieu. 

Adr.  Juss. 

Adrien  Jussieu,  son. 

Hagenb.      Hagenbach. 

Hall.           Haller. 

Ham.          Hamilton. 
Hanb.         Hanbury. 
Hanst.        Hanstein. 
Hartm.        Hartmann. 
Hartw.        Hartweg. 
Harv.          Harvey. 
Hass.          Hassall. 
Hassk.        Hasskarl. 

Kcemp. 
Karst. 
Kaulf. 
Kindb. 
Kirschl. 
Kit. 
Kcdr. 
Korth. 

Kaempfer. 
Karsten. 
Kaulf  uss. 
Kindberg. 
Kirschleger. 
Kitaibel.  ' 
Koelreuter, 
Korthals. 

Hausm.       Hausmann. 

Kostel. 
Kremp. 

Kosteletzky. 
Krempelhuber. 

Hebens.       Hebenstreit. 
Hedw.         Hedwig. 

Kromb. 

Kuetz. 

Krombholz. 
Kuetzing. 

Hegelm.       Hegelmaier. 

Hegetsch.     Hegetschweiler. 

L. 

Linnaeus. 

Heist.          Heister. 

Labill. 

La  Billardiere 

Heldr.         Heldreich. 

LcBSt. 

Laestadius. 

Helw.          Helwing. 

Lag. 

Lagasca. 

Hemsl.         Hemsley. 

LaU. 

Lallement.           [Marck) 

Henck.        Henckel. 

Lam. 

Lamarck  (Monnet  de  La 

Henfr.         Henfrey. 

Lamb. 

Lambert. 

Hensl.         Henslow. 

Lamour. 

Lamouroux. 

Herb.          Herbert. 

Langsd. 

Langsdorf. 

Herm.         Hermann. 

La  Peyr. 

La  Peyrouse. 

Hild.           Hildebrand. 

La  Pyl 

La  Pylaie. 

Hochst.        Hochstetter. 

Ledeb. 

Ledebour. 

Hoffm.        G.  F.  Hoffmann. 

Lehm. 

Lehmann. 

H.  Hoffm.  Hermann  Hoffmann 

Lem. 

Lemaire. 

/7q/7*/nanns.Hoffmannsegg. 

Lesq. 

Lesquereux. 

Hofm.         Hofmeister 

Less. 

Lessing. 

Hohen.        Hohenacker. 

Lestib. 

Lestiboudoia. 

Holmsk.      Holmskiold. 

Lto. 

Leveille'. 

Homb.         Hombron. 

L'Her. 

L'Heritier. 

Hook.         Win.  J.  Hooker. 

L'Herm. 

L'Henninier. 

Hook.f.      J.  D.  Hooker,  son. 

Liebm. 

Liebmann. 

Hopk.         Hopkirk. 

Light/. 

Lightfoot. 

Hornem.      Hornemann. 

Lilij. 

Lilijeblad. 

Hornsch.      Hornschuch. 

Lindb. 

Lindberg. 

Horsf.         Horsfield. 

Lindbl. 

Lindblom. 

Houst.         Houston. 

Lindenb. 

Lindenberg. 

Houtt.         Houttuyn. 

Lindh. 

Lindheimer. 

Huds.         Hudson. 

Lindl 

Lindley. 

Hueb.          Huebener. 

Linn. 

Linnaeus.    Also  £. 

Humb.         Humboldt. 

Linn.f. 

C.  Linnaaus,  son. 

J7RJT       5  Humboldt,  Bonpland.  and 

Lodd. 

Loddiges. 

'     -)      Kunth. 

Laefl. 

Loefling. 

388 


ABBREVIATIONS. 


Lees.    =    Loeselius. 

Naud.    =  Naudin. 

Lois.           Loiseleur-Delongschamps. 

Neck.          Necker. 

Loud.          Loudon. 
Lour.           Loureiro. 

Nee,  :  or    )  c  F  Neeg  VOQ  Esenbeck> 
N.abE.  J 

Ludw.         Ludwig. 

T.  Nees     T.F.L.  Nees  von  Esenbeck. 

Lumn.         Lumnitzer. 

Nestl.          Nestler. 

Lyngb.         Lyngbye. 

Newb.         Newberry. 

Newm.        Newman. 

Macf.          Macfadyen. 

Ncegg.         Noeggerath. 

Macgil.       MacGillivray. 

Nois.          Noisette. 

Magn.         Magnol.                     [stein. 

Nord.         Nordstedt. 

M.  Bfeb.     Marschall    von     Bieber- 

Not.            Notaris. 

Marsh.        Humphrey  Marshall. 

Nutt.           Nuttall. 

Mars.          Marsili. 

Nyl.           Nylander. 

Mart.          Martius. 

Nym.          Nyman. 

Mass.          Massalongo. 

Mast.          Masters. 

(Ed.           CEder. 

Maxim.       Maximowicz. 

(Erst.          CErsted. 

Med.           Medikus  or  Medicus. 

Oliv.           Olivier. 

Mean.      I  Meisner  or  Meissner. 
Meissn.     \ 

D.  Oliv.      D.  Oliver. 
Orb.            A.  or  C.  d'Orbigny. 

Meneg.        Menegliini. 

Orph.          Orphanides. 

Menz.          Menzies. 

Ort.             Ortega. 

Mert.           Mertens. 

Oudem.       Oudemans. 

Metten.        Mettenius. 

Mick.          Micheli. 

P.rfeBeauy.Palisot  de  Beauvois. 

Mirhr           ) 

j}j  n-n  r.      i  j^nfag  Michaux. 

Pall.           Pallas. 

MX.           ) 

Panz.          Panzer. 

Michx.  f.    F.  A.  Michaux,  son. 

Park.          Parkinson. 

Midden.      Middendorff. 

Parl.           Parlatore. 

Mill.           Philip  Miller. 

Pasq.          Pasquale. 

Mitt.  J.       John  S.  Mueller  or  Miller. 

Pav.            Pavon. 

Miq.             Miquel. 

Perl.           Perleb. 

Mirb.          Mirbel. 

Pers.           Persoon. 

Mitch.         John  Mitchell. 

Philib.         Philibert. 

Mitt.           Mitten. 

Planch.       J.  E.  Planchon. 

Moc.           Mo9ino. 

G.  Planch.  Gustave  Planchon. 

Molt.          Molkenboer. 

Pluk.           Plukenet. 

Mont.          C.  Montague. 

Plum.         Plumier,  Lat.  Plumerius. 

Moq.           Moquin-Tandon. 

Pcepp.         Poeppig. 

Moric.         Moricand. 

Pair.           Poiret. 

Moris.         Morison. 

Poit.           Poiteau. 

Morr.         Morren. 

Poll.            Pollich. 

Moug.         Mougeot. 

Post.           Postels. 

Muell.  Arg.  J.  Mueller  of  Argau. 

Pourr.         Pourret. 

F.  Muell.    Ferdinand  Mueller. 

Pringsh.      Pringsheim. 

0.  Muell.    Otto  Mueller  of  Denmark. 

Pritz.           Pritzel. 

Mukl.          Muhlenberg. 

Putter.        Putterlich. 

Munt           Munting. 

Murr.          J  A.  Murray. 

Rabenh.       Rabenhorst. 

A.  Murr      Andrew  Murray. 

Radlk.        Radlkofer. 

Raf.            Rafinesque-Schmaltz. 

Nacc.          Naccari. 

Rasp.          Raspail. 

Nag.           Naegeli. 

Red.            Redout^. 

ABBREVIATIONS. 


389 


Reich.    =  Reichard. 

Scop.    =    Scopoli. 

Reichenb.    H.  G.  L.  Reichenbach. 

Seem.         Seemann. 

Reichenb.f.  H.  G.  Reichenbach,  son. 

Sendt.         Sendtner. 

Reinw.         Reinwardt. 

Seneb.         Senebier. 

Reiss.          Reisseck. 

Ser.            Seringe. 

Retz.           Retzius. 

Seub.           Seubert. 

Rent.            Reuter. 

Sibth.          Sibthorp. 

Rich.           L.  C.  Richard. 

Sieb.           Sieber. 

fM.    JAchille  Richard. 

Sub.           Siebold. 
Soland.       Solander. 

Richards.    John  Richardson. 

Sow.           Sowerby. 

Richt.          Richter. 

Spenn.         Spenner. 

Ridd.          Riddell. 

Spreng.        SprengeL 

Riv.            Rivinus. 

Sternb.        Sternberg. 

Rachl.           Roehling. 

Steud.         SteudeL 

Raem.          J.  J.  Roemer. 

Stev.           Steven. 

M.  J.  RcemM.  J.  Roemer. 

Sull.           Sullivant 

^fJJ'  ^    |  Roemer  &  Schultes. 

Sw.            Swartz. 

Rap.           Roeper. 

Targ.          Targioni-TozettL 

Rohrb.         Rohrbach. 

Ten.            Tenore. 

Rostk.         Rostkovius. 

Thorns.        Thomas  Thomson. 

Rothr.         Rothrock. 

Thuitt.        Thuillier. 

Rottb.          Rottboell. 

Thunb.        Thunberg. 

Rotd.          Rottler. 

Thurb.         Thurber. 

Roum.         Roumegere. 

Thurm.       Thurman. 

Roxb.          Roxburgh. 

Tod.            Todaro. 

Roy.            Royen. 

Ton.          Torrey. 

Rudb.         Rudbeck. 

Ton.Sf  Gr.Torrey  &  A,  Gray. 

Rupr.          Ruprecht. 

Tourn.         Tournefort. 

Trait.          Trattinick. 

Sacc.          Saccardo. 

Trout.        Trautvetter. 

SadL          Sadler. 

Trev.           Treviranus. 

St.  EH.       A.  Sain^Hilaire. 

Trin.           Trinius. 

Salisb.         Salisbury. 

Tuck.         Tuckerman. 

Salm-Dyck.  Prince   Jos.   Salm-Riffer- 

Turcz.        Turczaninow. 

Sauss.         Saussure.     [schied-Dyck. 

Turn.          Turner. 

Schimp.       Schimper. 

Turp.          Turpin. 

Schk.           Schkuhr. 

Schlecht.      Schlechtendal. 

Vaitt.          Vaillant. 

Schleich.      Schleicher. 

Veill.          Veillard  or  Viefflard. 

Schomb.       Schomburgh. 

Vouch.        Vaucher. 

Schrad.       Schrader. 

Vent.           Ventenat. 

Schreb.        Schreber. 

ViU.            Villars,  or  VUlar. 

Schueb.        Schuebeler. 

Vis.             Visiani. 

Schult.         Schultes. 

Vittad.        Vittadini. 

Schultz    )  C.  H.  Schultz,  Bipontinus 

Viv.            Viviani. 

Bip.       J  (Zweibrucken). 

Vog.            T.  Vogel. 

Schum.        Schumacher. 

Schnitzl.      Schnitzlein. 

Wahl.         Wahlenberg. 

Schwcegr.     Schwaegrichea 

Wahlst.       Wahlstedt. 

Schwein.      Schweinitz. 

Wolds.       Waldstein. 

Schweinf.    Schweinfurth. 

Wall.          Wallich. 

Schwend.    Schwendener. 

WaUm.       Wallman. 

390 


ABBREVIATIONS. 


Wattr.  =  Wallroth. 


Walpere. 

Walter. 

Wangenheim. 

Warming. 

P.  W.  Watson. 
B.C.  Wats.H.  C.  Watson. 
S.  Wats.  Sereno  Watson. 

Weber. 

Weddell. 

Weinmann. 

Welwitsch. 

Wenderoth. 

Wendland. 
Wiks.         Wikstrom. 


Walp. 

Walt. 

Wang. 

Warm. 

Wats. 


Web. 

Wedd. 

Weinm. 

Welw. 

Wender. 


WUdb.  = 

Wildbrand. 

Willd. 

Willdenow. 

Willk. 

Willkomm. 

Wils. 

Wilson. 

Wimm. 

Wimmer. 

Wisliz. 

Wislizenus. 

With. 

Withering. 

Woodv. 

Woodville. 

Wulf. 

Wulfen. 

Zanard. 

Zanardini. 

Zetterst. 

Zetterstedt. 

Zucc. 

Zuccarini. 

Zuccag. 

Zuccagini. 

2.   ABBREVIATIONS  OF  NAMES  OF  ORGANS  AND  TERMS 
USED  IN  BOTANICAL  WRITINGS. 


JEst.  ^Estate,  in  summer. 

jEst.   ^Estivation. 

Alb.   Albumen. 

Anth.   Anther. 

Art.    Artificial. 

Auct.,  Auctt.  Auctorum,  of  authors. 

Aut.   Autumnal. 

B.  or  Beat.  Beatus,  "  the  late,"  re- 
cently deceased. 

Br.   Bract. 

Col.  Calyx. 

Cel.  Celeberrimus,  or  Very  cele- 
brated. 

Cent.   Centimetre. 

Cl.   Clarissimus. 

Char.   Character. 

Coll.  Collection. 

Cor.   Corolla. 

Cult.   Cultivated. 

Decim.  or  Dec™.  Decimetre. 

Descr.   Description. 

Diff.  Differentiae,  the  distinguishing 
marks. 

Ed.  Edition. 

Embr.   Embryo. 

Ess.  Essential,  as  Char.  Ess. 

Exd.   Excluding,  or  being  excluded. 

Excl.  Syn.  Excluding  the  synonym 
or  synonyms. 


Fam.   Family. 

Fil.  Filament  of  the  stamen. 

Fl.  Flower  (flos) ;  Flora,  or  some- 
times Floret,  it  flowers. 

Fcem.   Female  plant,  flower,  &c. 

Fol.  Folium,  leaf. 

Fr.  Fruit. 

Fructif.   Fructification. 

Gen.   Genus  or  Generic. 

Germ.  Germen,  Linnaean  name  for 
ovary ;  also  Germination. 

H.   Herbarium. 

Hob.  Habitat,  place  of  growth; 
sometimes  for  Habeo,  I  have. 

Herb.   Herbarium. 

Hart.   Hortus,  garden. 

Horttd.  Hortulanorum,  of  the  gar- 
deners. 

Ic.   Icon,  a  plate  or  figure. 

III.   Illustris,  illustrious. 

Ined.   Unpublished. 

Inf.    Inferior. 

Infl.   Inflorescence. 

Inv.   Involucre. 

Lot.   Lateral,  or  relating  to  width. 

Lin.  Linea,  a  line  (the  12th  of  an 
inch). 

Lit.,  Litt.  In  a  letter  or  letters. 

/.  c.  Loco  citato,  in  the  place  cited. 


ABBREVIATIONS. 


391 


Masc.   Male  plant,  flower,  &c, 

Mill,   or  mm.   Millimetre. 

Mss.   Manuscripts. 

Mus.  Museum. 

N.  or  No.  Number. 

Nat.  Natural. 

Nom.  Nomen,  name. 

Obs.   Observation. 

Ord.   Order. 

Ov.   Ovary. 

p.  Page,  or  sometimes  Part. 

Ped.  Peduncle    or  Pedicel,   or  for 

Pedalis,  a  foot  long  or  high. 
Peric.   Pericarp. 
Perig.   Perigonium. 
Pet.   Petal  or  Petiole. 
Pist.   Pistil. 
Plac.   Placenta. 
Poll.    Pollicaris,  an  inch  long. 
p.  p.  Pro  parte,  in  part. 
Prodr.  or  Prod.   Prodromus. 
Rad.   Radix,  root ;  or  Radical. 
Ram.   Ramus,  branch. 
«.   Seu,  or  Sive,  Latin  for  or. 
Sect.   Section. 
Segm.   Segment. 
Sem.  Semen,  seed. 


Sep.   Sepal. 

Ser.   Series. 

-Sice.  Siccatus  or  Siccus,  dried  or  dry. 

Spec,  or  Sp.   Species,  or  specimen. 

Spont.   Spontaneous. 

Stam.  Stamen  or  Staminate. 

Sup.   Superior. 

Syn.   Synonym  or  Synopsis. 

T.  or  Tab.   Tabula,  plate. 

T.  Toiuus,  volume. 

V.  Volume  :  sometimes  for  Vel,  or ; 
sometimes  Vide,  see. 

Var.    Variety. 

Veg.   Vegetation,  characters  of 

Fern.   Vernal. 

v.  s.   Visa  sicca,  or  Vidi  siccam. 

v.  v.  Visa  viva,  or  Vidi  vivam ;  the 
first  indicating  that  a  dried  speci- 
men of  the  plant,  the  second  that 
the  living  plant  has  been  exam- 
ined. 

v.  s.  c.  and  v.  s.  s.,  indicates  that 
the  dried  specimen  was  cultivated 
(c)  or  spontaneous  (s). 

v.  v.  c.  and  v.  v.  s.,  that  the  living 
plant  seen  was  cultivated  (c)  or 
spontaneous  (s). 


SIGNS. 


1.  SIGNS  USED  BY  LINKEUS. 

0  An  annual  plant. 

$  A  biennial. 

y.  A  perennial. 

5  A  tree  or  shrub. 

#  Affixed  to  a  reference,  means  that  a  good  description  will  be  found  there, 

t  Indicates  an  obscure  or  doubtful  species. 


2.    SIGNS  USED   BY  DECANDOLLE  AND   LATER 
WRITERS. 

O    A  monocarpic  plant,  t.  e.  which  dies  after  once  flowering  and  fruiting, 

either  annual  or  biennial,  or  of  longer  duration. 
©   Annual. 

Biennial. 

Monocarpic  perennial,  such  as  Agave. 


392  SIGNS. 

"21    Perennial  herb. 

*)    Suffrutex,  an  undershrub. 

*>     Frutex,  a  shrub. 

5    Arbuscula,  a  tree-like  shrub  of  ten  to  twenty-five  feet  in  height 

5     Arbor,  a  tree. 

r\   A  climbing  plant. 

A    An  evergreen. 

$    Male  plant  or  flower. 

9    Female  plant  or  flower. 

"5*   Hermaphrodite  plant  or  flower. 

00    Indefinitely   numerous,  e.  g.  co-andra,  polyandrous. 

?  A  sign  of  doubt.  "  Thalictrum  ?  Japonicum,"  doubts  if  the  plant  is 
really  a  Thalictrum.  "  Thalictrum  Japonicum,  Thunb.?  "  doubts  if  the 
plant  in  hand  is  truly  the  species  of  Thunberg.  Thalictrum  Japoni- 
cum, Thunb.,  Willd.?  doubts  whether  Willdenow's  T.  Japonicum  is 
really  that  of  Thunberg. 

1  A  sign  of  certainty.  As  "  Thalictrum  anemonoides,  Michx. !  Fl.  Bor.  Am. 
p.  322,"  as  used  by  DeCandolle,  affirms  that  he  has  seen  an  authentic 
original  specimen  of  this  author.  Affixed  to  the  name  of  a  collector, 
as  "  Virginia,  Clayton  !  "  it  affirms  that  the  writer  has  examined  a 
specimen  collected  by  the  person  to  whose  name  it  is  appended. 

-   Between  two  figures,  as  in  "  Stamens  6-10,"  indicates  the  extremes  of 

difference,  as  that  the  stamens  are  from  five  to  ten. 

0  ' "  The  signs  for  degrees,  minutes,  and  seconds,  as  1°,  2',  3",  are  used  in 
Gray's  Manual  of  Botany  of  the  Northern  United  States,  for  feet  (°), 
inches  ('),  and  lines  (").     With  European  authors,  usually  the  sign 
for  minutes  is  for  feet ;  that  of  seconds  for  inches  :  thus  V,  a  foot 
high  ;  1",  an  inch  long ;  and  I'",  a  line  long. 
O=  Cotyledons  accumbent  to  the  radicle. 
O II   Cotyledons  incumbent  on  the  radicle. 


GLOSSARY, 

OR 

DICTIONARY    OF    BOTANICAL    TERMS,   ENGLISH 
AND    LATIN, 

COMBINED    WITH    AN    INDEX. 


THIS  Glossary  is  intended  to  contain  all  the  principal  technical  terms  (substan- 
tive as  well  as  adjective)  of  structural  and  systematic  Botany,  as  far  at  least  as 
concerns  Phsenogamous  plants.  Most  of  the  special  terms  relating  to  the  lower 
Cryptogamia  and  to  Vegetable  Anatomy  and  Physiology  are  relegated  to  the  vol- 
umes devoted  to  those  departments.  The  annexed  numbers  refer  to  pages  of  this 
volume.  Very  many  of  the  terms  are  seldom  employed,  or  are  wholly  out  of  use. 
The  principal  Latin  terms  are  given  separately  only  when  there  is  no  English  equiv- 
alent differing  merely  in  the  termination.  When  the  word  is  essentially  the  same, 
the  Latin  termination  (of  adjectives  in  the  nominative  masculine  only)  is  annexed 
to  the  English  word  in  a  parenthesis.  The  changed  termination  goes  back  mostly 
to  the  penultimate  consonant.  It  is  unnecessary  in  a  work  like  this  to  accentuate 
all  the  technical  words;  but,  in  the  case  of  words  liable  to  mispronunciation,  an 
accent-mark  is  placed  over  the  syllable  which  takes  the  principal  accent.  The 
glossary,  as  here  drawn  up,  may  serve  to  indicate  the  meaning  of  the  commoner 
descriptive  specific  names  of  plants. 


A,  privative,  as  the  initial  in  many 
words  of  Greek  derivation,  signifies 
the  absence  of  the  organ  mentioned; 
as,  ^Ipetalous,  destitute  of  petals; 
Aphyllous,  leafless.  In  words  be- 
ginning with  a  vowel,  this  prefix  is 
changed  to  an;  as,  .4raanthous,  flow- 
erless  ;  .4«antherous,  antherless. 

Abbreviations,  385. 

Aberrant.  Wandering,  applied  to  spe- 
cies, genera,  &c.,  which  differ  in  some 
respect  from  the  usual  or  normal  char- 
acter of  the  group  they  belong  to. 

Abnormal  (Abnormis).  Differing  from 
the  normal  or  usual  structure. 

Aboriginal.    Strictly  native ;  indigenous. 

Abortion  (Abortus).  Imperfect  develop- 
ment or  non-development  of  an  organ  ; 
179,  187. 

Abortive  (-ivus).    Defective  or  barren. 

Abrupt  (Abruptus).  Terminating  sud- 
denly ;  the  opposite  of  tapering. 


Abruptly  pinnate.  Pinnate  without  a 
terminal  leaflet  or  appendage  ;  101. 

Acantkocladous  (-us).  Having  spiny 
branches. 

Acanthqphorous  (-us).     Spine-bearing. 

Acaulescent  (-ens).  Stemless,  or  appar- 
ently so,  with  no  proper  caulis ;  45. 

Acaulis.  Stemless ;  same  as  Acaulescent. 

Accessory.  Something  additional,  or  of 
the  nature  of  appendage. 

Accessory  Suds,  44. 

Accessory  Fruits,  300. 

Accrescent  (-ens).  Increasing  in  size 
with  age,  as  often  occurs  with  the 
calyx  after  flowering. 

Accrete  (-us).  Grown  together,  or  con- 
solidated with  some  contiguous  body. 

Accumbent  (-ens).  Lying  against  an- 
other body. 

Accumbent  Cotyledons.  With  edges 
against  the  radicle;  313. 

Acephalous  (-us).    Headless- 


394 


GLOSSARY. 


jicerose  (-<fow).  Needle-shaped,  like  the 
leaves  of  Pines. 

Acetdbuliform  (-ormis).  In  the  form  of 
a  shallow  open  cup  or  saucer. 

Ackcenium  or  Achenium.  A  small,  dry 
and  hard,  one-celled,  one-seeded,  inde- 
hiscent  fruit;  strictly  one  of  a  single 
and  free  carpel ;  but  extended  to  simi- 
lar ones  of  more  than  one  carpel,  and 
also  with  adnate  calyx ;  294.  (Achee- 
nium  is  etymologically  the  proper 
orthography ;  but  achenium  is  be- 
coming the  commoner  form.) 

Achcenocarp  (-arpium).  General  name 
of  a  dry  and  indehiscent  fruit ;  292. 

Achenodium.  Such  a  double  achenium 
as  that  of  Umbelliferse ;  a  Cremo- 
carp. 

Achlamydeous  (-ens).  Destitute  of  peri- 
anth; 191. 

Acicula.    A  bristle. 

Acicular  (-am).  Bristle-shaped,  or  slen- 
der needle-shaped. 

Acindciform  (-ormis).  Scymitar-shaped ; 
curved  with  rounded  point,  thicker  on 
the  straighter  edge  than  on  the  con- 
vex edge. 

Acinows.     Like  grapes  or  grape-seed. 

Acinus.  Classically  a  berry,  particu- 
larly a  grape,  or  its  stony  seed,  or  a 
bunch  of  berries ;  now  sometimes  ap- 
plied to  the  separate  carpels  of  an 
aggregate  baccate  fruit,  or  to  the  con- 
tained stone  or  seed ;  297. 

Acorn.     Fruit  of  the  Oak. 

Acotyledon,  pi.  Acotyledons,  Acotyledones. 
A  plant  or  plants  destitute  of  coty- 
ledon, or 

Acotyledonous  (-eus).  Without  cotyle- 
dons ;  as  the  embryo  of  Cuscuta ;  26, 
38.  Mostly  applied,  as  by  Jussieu, 
to  plants  which  have  no  proper  seed 
nor  embryo,  and  therefore  no  cotyle- 
don ;  339. 

Acramphibrya.  Plants  producing  side  as 
well  as  terminal  buds  or  growths  ;  341. 

Acrdbrya.  Plants  growing  from  apex 
only;  341. 

Acrogen  (Acrdgence).  Name  of  class  of 
plants  which  in  growth  are  said  to  be 

Acroyenous.  Growing  from  the  apex 
or  by  terminal  buds  only. 

Acrosdrcum.  Desvaux's  name  for  a 
berry  from  an  ovary  with  adnate 
calyx. 

Acrotpira.  An  old  name  of  the  plu- 
mule of  a  grain  in  germination. 

Aculeate  (-eatus).  Prickly;  beset  with 
aculei. 


Aculeosus.    Abounding  with  prickles. 

Aculeolate  (-atus).  Beset  with  diminu- 
tive prickles,  or 

Aculeoli.    Diminutive  of  aculei. 

Aculeus.  A  prickle;  a  pointed  small 
excrescence  of  the  bark. 

Acumen.    A  tapering  point. 

Acuminate  (-atus).  Ending  in  a  tapering 
point;  96. 

Acute  (Acutus).  Terminating  in  an  acute 
angle;  97. 

Acropetal.  Developing  from  below  up- 
ward, or  from  base  toward  apex. 

Actinomorphous  (-us).  Capable  of  bi- 
section through  two  or  more  planes 
into  similar  halves,  as  is  a  regular 
symmetrical  flower;  175. 

Acutiusculus.    Somewhat  acute ;  acutish. 

Adelphous  (-us,  Adelphi,  brothers).  Sta- 
mens with  coalescent  or  clustered  fila- 
ments are  monadelphous,  diadelphous, 
&c.,  according  to  the  number  of  Add- 
phia  or  brotherhoods. 

Aden.  Greek  for  gland,  is  compounded 
with  Greek  words  with  this  meaning; 
as,  Adenophorus,  gland-bearing;  Ad- 
enophyllus,  leaves  bearing  glands,  &c. 

Adylutinate  (-atus).     Same  as  accrete. 

Adherent  (Adhcerens).  Generally  same 
as  adnate ;  may  refer  to  adhesions  not 
congenital. 

Adnate,  (-atus).  Congenitally  united  to  ; 
as  the  calyx-tube  of  the  gooseberry  to 
the  ovary;  182.  Adnate  anther  is  one 
seemingly  borne  on  the  outer  or  inner 
face  of  the  filament ;  i.  e.  extrorsely  or 
introrsely  fixed  by  its  whole  length  to 
the  connective ;  253. 

Adnation.  The  state  of  being  adnate; 
179,  181. 

Adpressus.     Latin  of  appressed. 

Adscendens.     Latin  of  ascending. 

Adsurgens.     Latin  of  assurgent. 

Adventitious,  Adventive.  That  which  has 
come  from  abroad  or  as  a  stranger ;  as 
a  plant  lately  or  by  chance  introduced 
from  another  country. 

Adventitious  Buds ;  45. 

jEquilaterdlis.    Equilateral,  equal-sided. 

jEqualiftorus.  When  all  the  flowers  of 
the  same  head  or  cluster  are  alike  in 
form  as  well  as  character. 

^(judlis,  sEquans.     Equal ;  equalling. 

Aerial  roots,  &c.  ;  33. 

Aerophytes.     Air-plants;  35. 

jEruginosus.     Verdigris-colored. 

jEstival  (-dlis).     Relating  to  summer. 

^Estivation  (-io).  The  disposition  of  the 
parts  of  a  flower  in  the  bud ;  132. 


GLOSSAEY. 


395 


^Eterio.  A  form  of  aggregate  fruit;  300. 
^Etheogamia,  ^Ethogamous,  340. 
Affinity.     True   and   near   relationship; 

m,  330. 
Agamous or  Agamic.   Destitute  of  sexes. 

Agglomerate.         (  HeaPed  or  clTded 

Aggregate^),     mto  a  dense  cluster, 
I  but  not  cohering. 

Aggregate  Fruits.  Those  formed  of 
aggregate  carpels  of  the  same  flower; 
298,  301. 

Agrestis.     Growing  in  fields. 

Air-plants.  Plants  unconnected  with  the 
ground;  35. 

Akene,  Akenium.    See  Achsenium. 

Ala  (pi.  alee).  A  wing.  Also  the  side 
petals  of  a  papilionaceous  corolla; 
185.  Has  also  been  used  in  the  sense 
of  axilla. 

Alabdstrum.    A  flower-bud ;  40. 

Alar  (Alaris).  From  ala  in  the  sense  of 
axilla,  therefore  axillary  or  in  the  forks. 

Alate  (-atus).     Winged. 

Albescens,  Albicans.  Whitened,  whitish, 
or  hoary. 

Albumen  of  the  seed.  Any  deposit  of 
nutritive  material  within  the  seed- 
coats,  and  not  in  the  embryo ;  14,  309. 

Albuminous  or  Albuminose  (Albuminosus). 
Said  of  seeds  provided  with  albumen ; 
13,  309. 

Alburnum.  Sapwood;  the  newer  wood 
of  an  exogenous  stem ;  80. 

Albus.     White. 

Allagostemonous.  With  stamens  alterna- 
tively inserted  on  the  torus  and  on  the 
petals. 

Alliaceous  (-eus).  Having  the  smell  of 
garlic. 

Alliance.     Synonym  of  Cohort;  326. 

Allogamy.  Fecundation  of  the  ovules  of 
a  flower  by  other  than  its  own  pollen ; 
cross  fertilization,  216. 

Alpestrine  (Alpestris).  Growing  on 
mountains  below  an  alpine  region 
or  one  unwooded  from  cold. 

Alpine  (-inus).  Growing  on  the  higher 
parts  of  the  Alps,  or  (by  extension  of 
meaning)  on  other  mountains  above 
the  limits  of  trees. 

Alternate  (Altirnus).  One  after  an- 
other; as  of  leaves  placed  singly 
instead  of  in  pairs  (opposite)  or  in 
whorls.  Also,  standing  before  inter- 
vals ;  as  stamens  alternate  with  petals 
instead  of  before  them;  6,  119. 
Alternative  (-ivus).  In  aestivation,  with 
an  inner  whorl  alternating  with  an 
outer  one;  134,  136. 


Alv^olate(-atus).  Honeycombed;  having 
deep  angular  cavities  (Alveoli)  sepa- 
rated by  thin  partitions,  as  the  recep- 
tacle of  cotton-thistle. 

Ambitus.  The  ray  or  circumference  of 
a  head,  &c. 

Ament  (Amentum).  A  catkin,  or  pe- 
culiar scaly  spike ;  150. 

Amentaceous  (-eus).  Bearing  catkins,  or 
catkin-like. 

Amorphous  (-us).  Shapeless;  of  in- 
definite form. 

Amphanthium.  One  of  the  (needless) 
names  coined  for  a  dilated  receptacle 
of  inflorescence. 

Amphibrya.  Equivalent  to  Monocotyle- 
dones;  341. 

Amphicarpous  (-us).  Producing  two 
kinds  of  fruit. 

Amphigamous  Cryptogams,  340. 

Amphigastria.  Peculiar  leaves  (of  He-  > 
paticse)  imitating  stipules. 

Amphisarca.  A  hard-rinded  berry,  or 
fruit  succulent  within  and  woody  or 
crustaceous  without,  as  a  calibash. 

Amphispermium.  Link's  name  for  a 
one-seeded  pericarp  which  is  con- 
formed to  the  seed ;  an  akene. 

Amphitropous  (-us),  wrongly  Amphi- 
tropal.  Turned  both  ways;  applied 
to  an  ovule  with  hilum  intermediate 
between  micropyle  and  chalaza ;  279. 

Amphora.  A  pitcher;  and  the  lower 
part  of  a  pyxis. 

Amplectens,  Amplexans,  Amplexus.  Em- 
bracing, clasping. 

Amplexicaul  (-aulis).  Clasping  a  stem, 
as  does  the  base  of  certain  leaves. 

Ampliate  (-atus).    Enlarged  or  dilated. 

Ampulla.  A  bladder  or  flask-shaped 
organ,  as  of  Utricularia. 

Ampullaceous  (-us),  or  Amputtaform. 
In  the  form  of  a  bladder  or  short  flask. 

Amylaceous  (-eus).  Resembling  or  com- 
posed of  starch,  or  Amylum. 

Amyloid.     Analogous  to  starch. 

Analogy  (Analogia).  Likeness  in  cer- 
tain respects.  As  distinguished  from 
affinity,  it  means  resemblance  in  cer- 
tain respects  only,  not  in  the  plan  of 
structure.  Thus,  a  Ranunculus  is 
analogous  to  a  Potentilla,  but  there 
is  no  near  affinity  or  relationship  be- 
tween the  two.  And  the  tendril  of  a 
Pea,  that  of  a  Smilax,  and  that  of 
the  Grape-vine  are  analogues ;  i.  e.,  are 
analogous  organs,  but  are  not  homo- 
logues ;  for  the  first  answers  to  a  leaf, 
the  second  to  stipules,  and  the  third 


396 


GLOSSARY. 


to  a  stem.  The  spur  of  a  Larkspur  is 
analogous  to  one  of  the  five  spurs  of 
Columbine,  but  not  homologous  with 
it;  for  the  first  is  a  sepal,  and  the 
second  a  petal. 

Anandrous.     Destitute  of  stamens. 

Anantherous  (-MS).    Destitute  of  anthers. 

Ananthous  (-us).    Flowerless. 

Anastomdsis.  The  connection  of  veins, 
&c.,  by  cross-veins,  forming  reticu- 
lation. 

Andtropous  (-««),  wrongly  Anatropal. 
The  reversed  ovule,  with  micropyle 
close  by  the  side  of  the  hilum,  and 
chalaza  at  the  opposite  end ;  279. 

Anceps,  Engl.  Ancipital.  Botanically 
always  used  in  the  sense  of  two- 
edged. 

Ander,  andra,  andrum.  In  Greek  com- 
pounds, the  male. 

Andro-dicecious.  With  flowers  on  one 
plant  hermaphrodite,  and  on  another 
staminate  only;  191. 

Andrcedum.  The  stamens  of  a  flower 
collectively ;  165,  249. 

Andrdgynous  (-us).  Said  of  an  inflores- 
cence composed  of  both  male  and 
female  flowers. 

jindrophore  (Andr6phorum).  A  sup- 
port or  column  on  which  stamens  are 
raised. 

Androus.     See  Ander. 

Anemdphilous.  Literally  wind-loving. 
Said  of  flowers  which  are  fecundated 
by  wind-borne  pollen ;  217. 

Anfractuosus.  Abruptly  bent  hither  and 
thither,  as  the  stamens  of  Cucur- 
bita. 

Angiocarpous  (-«*).  When  a  fruit  is 
covered  by  some  envelope. 

Angiospermia.  A  Linmean  artificial 
order;  337. 

Anffiospermous,  Angiospermce,  Angio- 
sperms.  Plants  with  seeds  borne  in  a 
pericarp;  259. 

Angular  Divergence  of  leaves ;  123. 

Anisomerous  (-us).  Unequal  in  number 
in  the  different  circles  of  the  flower; 
unsymmetrical. 

Anisopetalous  (-us).  With  unequal  petals. 

Anisophyllous  (-us).  Unequal-leaved; 
f.  e.,  the  two  leaves  of  a  pair  unequal. 

Anisostemonous  (-us).  When  the  sta- 
mens are  not  of  the  number  of  the 
pstals. 

Annotinus.  A  year  old,  or  in  yearly 
growths. 

Annual  (Annuus).  Of  only  one  year's 
duration;  30. 


Annular  (-arts).  In  the  form  of  a  ring; 
or  marked  transversely  by  rings. 
The  latter  more  properly. 

Annulate  (-atus).    Marked  with  rings. 

Annulus.  A  ring,  such  as  that  with 
which  the  sporangia  of  some  Ferns 
and  Mosses  are  furnished. 

Anophytes  (Anophyta).  Name  of  group 
comprising  Mosses,  &c. 

Anttposition  Same  as  Superposition; 
179,  195. 

Anterior,  as  to  position,  denotes  the  front 
side,  or  averse  from  the  axis  of  inflo- 
rescence; 160. 

Anthela.  A  deliquescent  and  paniculate 
cyme,  with  median  ramification,  and 
the  lateral  axes  overtopping  the 
central,  as  in  Juncus  tenuis,  &c. 
May  be  either  a  Drepanium  or  a  Rhi- 
pidium. 

Anthemy,  Anthemia.  A  flower-cluster 
of  any  kind ;  144. 

Anther  (Anthera).  The  polliuiferous 
part  of  a  stamen;  165,  251. 

AntherUKum.  An  analogue  of  the  an- 
ther in  Cryptogams. 

Antheriferous  (-«*•).     Anther-bearing 

Anthesis.  The  time  at  which  a  flower 
is  perfected  and  opens;  or  the  act  of 
expansion  of  a  flower. 

Anthocarpous(-us),Anfhoc(trpium.  Fruits 
in  which  some  organ  exterior  to  the 
pericarp  is  concerned  ;  300. 

Anthoclinium.  Name  of  a  receptacle  of 
inflorescence,  such  as  that  of  Com- 
positae. 

Anthodium.  A  name  for  the  head  of 
flowers  (or  so-called  compound  flowers) 
of  Composite;  147. 

Antholysis.  A  retrograde  metamor- 
phosis of  a  flower,  in  which  normally 
^  combined  parts  are  separated. 

Jlnthophore  (Anthdphorum).  The  stipe 
when  developed  between  calyx  and 
corolla;  212. 

Anthus  or  Anthos.  A  flower,  in  Greek 
compounds. 

Anticous  (Anticus).  Facing  anteriorly; 
253. 

Antitropous  (-us),  less  properly  Anttiro- 
pal.  Said  of  an  embryo  with  *adicle 
pointing  to  the  end  of  the  seed  oppo- 
site the  hilum;  312. 

Antrorse.     Directed  upward  or  forward. 

Apetalous  (-us).    Having  no  petals ;  190. 

Apex.  Besides  its  ordinary  meaning, 
the  top  of  a  thing,  it  was  once  the 
technical  name  of  an  anther;  166. 

Aphyllous  (-us).    Leafless. 


GLOSSABY. 


397 


Apical  (-alis).  Relating  to  the  apex  or  tip. 

Jlpices.  The  name  for  anthers  anterior 
to  Ludwig  and  Linnaeus ;  166. 

Apiculate  (-us).  Ending  in  a  short 
pointed  tip  or  apicula. 

Apocarpous  (-us).  When  carpels  of  a 
gyncecium  are  separate ;  261,  262. 

Apdphysis.  An  enlargement  or  swelling 
of  the  surface  of  an  organ  at  some  par- 
ticular part. 

Apothecia.  The  "shields  "  or  fructify- 
ing disks  of  Lichenes. 

Apotropous  (-us).  Said  of  an  anatropous 
ovule  which  when  pendulous  has 
rhaphe  averse ;  282. 

Appendage,  Appendix.  Any  superadded 
or  subsidiary  part. 

Appendiculate  (-atus).  Furnished  with  a 
small  appendage  (Appendiculum),  or 
with  any  appendage. 

Appositus.    Placed  side  by  side. 

Appressed  (Lat.  Adpressus).  Lying  flat 
against  or  together  for  the  whole 
length. 

Apricus.     Growing  in  dry  sunny  places. 

Apterous  (-us).     Wingless ;  not  alate. 

Aquatic  (-icus).     Living  in  water. 

Aqudtilis.     Living  under  water. 

Arachnoid  (-oideus).  Cobwebby  ;  com- 
posed of  slender  entangled  hairs. 

Araneose  (-osus),  Araneus.  Like  spider- 
web  ;  same  as  Arachnoid. 

Arbor.    A  tree  ;  50. 

Arboreous  (Arboreus).  Tree-like,  or  re- 
lating to  a  tree. 

Arborescent  (-ens).  Tree-like;  approach- 
ing the  size  of  a  tree. 

Arboretum,  also  Arbustum.  A  place 
where  trees  are  grown;  an  arranged 
collection  of  trees. 

Arbuscula.  A  small  shrub  of  tree-like 
growth  or  form. 

Arbuscularis.   Ramified  like  a  little  tree. 

Archegonium.  The  spore-case  of  mosses, 
^c.,  in  an  early  state. 

Arcuate  (-atus).  Moderately  curved,  as 
if  bent  like  a  bow. 

Areola,  pi.  Areolce.  Spaces  marked  out 
on  a  surface,  as  by  the  reticulation  of 
veins,  &c. 

Areolate  (-atus).    Marked  with  areolae. 

Arenosus,  Arenarius.  Growing  in  sand 
or  sandy  places. 

Argentate  (-atus,  Argenteus).  Silvery, 
or  shining  white  with  a  tinge  of  gray. 

Argillosus.     Growing  in  clayey  soil. 

Argos.  Greek  for  pure  white,  used  in 
compounds;  as,  argophyllus,  white- 
leaved. 


Argutus.  Sharp-toothed;  said  of  the 
serration  of  leaves. 

Argyros.  Greek  for  silvery;  used  in 
compounds;  as,argyrophyllus,  silvery- 
leaved. 

Arhizal  (Arhizus).     Rootless. 

Arillate  (-atus).     Having  an  arillus. 

Aril,  Arillus.  An  extraneous  or  late- 
formed  seed-coat  or  covering,  or  an 
appendage  growing  from  or  about  the 
hilum  of  a  seed;  308. 

Arilliform  (-ormis).  In  the  form  of  an 
arillus. 

Arillode,  Arillodium.  A  false  arillus, 
or  one  which  does  not  originate  from 
or  below  the  hilum,  but  from  the 
micropyle  or  rhaphe;  309. 

Arista.     An  awn. 

Aristate  (-atus).  Awned;  bearing  an 
arista. 

Aristulate  (-atus).  Bearing  a  diminu- 
tive awn. 

Arrect  (Arrectus).  Brought  into  an 
upright  position. 

Arrow-shaped,  Arrow-headed.  Same  as 
Sagittate;  96. 

Articulated  (-atus).  Jointed,  or  having 
the  appearance  of  a  joint  or  articula- 
tion (Articulus).  As  of  the  word  joint 
itself,  the  context  must  show  whether 
the  articulations  mean  the  portions 
which  are  connected  by  a  joint,  or  the 
place  of  connection. 

Artificial  Classification,  331. 

Ascending  (Adscendens).  Rising  upward. 
Sometimes  used  for  directed  upward, 
as  when  the  stem  is  termed  the  As- 
cending Axis  (11);  more  commonly 
denotes  curving  or  rising  obliquely 
upward;  53. 

Ascidium.  A  pitcher-shaped  or  flask- 
shaped  organ  or  appendage ;  111. 

Ascus.  A  sac ;  a  kind  of  spore-cases,  as 
in  certain  Fungi  and  Lichenes. 

Aspdragi,  A  name  for  Turiones,  or  any 
scaly  shoots  from  underground,  as 
those  of  Asparagus. 

Aspergilliform  (-ormis).  Brush-shaped, 
i.  e.  like  the  aspergiUum,  or  brush 
used  to  sprinkle  holy  water ;  made  up 
of  numerous  spreading  hairs,  &c.,  in 
a  tuft,  as  the  stigmas  of  Grasses. 

Asperous  (Asper).     Rough  to  the  touch. 

Assimilation.  The  action  or  process  by 
which  extraneous  matter  or  crude  food 
is  converted  into  vegetable  matter. 

Assurgent  (Adsurgens).  Rising  or  curv- 
ing upward ;  53. 

Astichous  (^ts).    Not  in  rows. 


398 


GLOSSARY. 


jistomotu  (-itf).  Without  a  stoma  or 
mouth. 

Atavism  (-mas).  Ancestral  resemblance. 

Ater.    Pure  black. 

Mhera.    Greek  for  Arista  or  Awn. 

Atrdtus.    Blackened  or  turning  black. 

Mropous  (-us),  wrongly  AtropaL  Not 
turned ;  applied  to  an  ovule  the  same 
as  orthotropous ;  277. 

Attenuate  (-atus).  Slenderly  tapering 
or  narrow. 

Auctus.  Same  as  accrescent;  enlarged 
after  flowering ;  augmented  by  an  ad- 
dition. 

Augmentation.  Increase  beyond  the 
normal  number;  179,  200. 

Aurantiacus.    Orange-colored. 

Auratus,  Aureus.  Golden-colored,  or 
yellow  with  golden  lustre. 

Auricle  (Auricula).  An  ear  or  ear- 
shaped  appendage. 

Auriculate  (-aim).  Furnished  with  an 
auricle;  96. 

Autocarpous.  A  fruit  consisting  of  peri- 
carp alone,  having  no  adnate  parts. 

Autogamy.  Close-fertilization,  the  fe- 
cundation of  a  flower  by  its  own  pol- 
len; 215,  216. 

Avenius.    Veinless. 

Awl-shaped.  Narrow,  terete  or  some- 
what so,  and  attenuate  from  a  broader 
base  to  a  slender  or  rigid  point 

Awn.  A  bristle-shaped  appendage,  such 
as  the  beard  of  Rye  and  Barley. 

Awned.     Furnished  with  an  awn. 

Axil  (Axilla).  The  angle  formed  on  the 
upper  side  of  the  attachment  of  a  leaf 
with  the  stem,  or  the  point  just  above 
this  attachment ;  6. 

Axillary  (-dris).  In  or  relating  to  an 
axil;  7. 

Axile,  Axial  (Axitis).  Relating  or  be- 
longing to  the  axis. 

Axis.  The  stem ;  the  central  part  or 
longitudinal  support  on  which  organs 
or  parts  are  arranged ;  the  central  line 
of  any  body. 

Bacca.    A  berry ;  299. 

Baccate  (-atus).  Berry-like;  pulpy 
throughout. 

Baccetum.  An  aggregation  of  berries 
in  one  flower ;  300. 

Badius.    Chestnut-brown. 

Balausta.  Name  applied  to  the  fruit 
of  the  Pomegranate,  with  firm  rind, 
crowned  with  the  lobes  of  an  adnate 
calyx,  baccate  within,  and  many- 
seeded. 


Banner.  The  vexillum,  standard,  or 
upper  petal  of  a  papilionaceous  co* 
rolla;  184. 

Barb.  A  bristle  or  stout  hair,  which  is 
hooked  or  double-hooked,  or  retrorsely 
appendaged  at  the  tip. 

Barba.    Beard. 

Barbate  (-atus).  Bearded;  beset  with 
long  and  weak  hairs. 

Barbellate  (-atus).  Beset  with  shorter 
and  stiffer  hairs  or  barbellce. 

Barbellulate  (-atus).  Diminutive  of  the 
preceding. 

Bark.  The  rind  or  cortical  portion  of  a 
stem,  especially  of  an  exogen ;  76. 

Basal  (Basilaris).    Relating  to  the  base. 

Basal-nerved.  With  nerves  all  from  the 
base  of  the  leaf;  92. 

Base  (Basis).  The  extremity  by  which 
an  organ  is  attached  to  its  support. 

Basidia.  Cells  of  the  fructification  of 
Mushrooms  which  bear  the  spores. 

Basifixed  (-us).  Attached  by  the  base 
or  lower  end ;  253. 

Basigynium.  Synonym  of  Carpophore 
or  Thecaphore. 

Basinerved  (-ius).  When  the  ribs  pro- 
ceed from  the  base  of  a  leaf. 

Basipetal.  Developing  from  apex  to- 
ward the  base. 

Bast,  or  Bass.    Inner  fibrous  bark ;  77. 

Bast-cells.  The  essential  components  of 
bast ;  long  and  flexible  but  thick-walled 
attenuated  cells;  77. 

Beak.    A  narrowed  or  prolonged  tip. 

Beaked.    Ending  in  a  beak. 

Bell-shaped.  Same  as  Campanulate ;  249. 

Berried.    Baccate. 

Berry.  A  fruit,  the  whole  pericarp  of 
which  is  fleshy  or  pulpy;  299. 

Bi-  or  Bis.  As  a  prefix  to  Latin  words 
(Greek  words  have  Di-),  two,  twice, 
or  doubly. 

Biacuminate  (-atus).  Two-pointed,  as 
malpighiaceous  hairs,  fixed  by  the 
middle  and  tapering  to  each  end. 

Biarticulate  (-atus).     Two-jointed. 

Biauriculate  (-atus).    Two-auricled. 

Bibracteate  (-atus).    With  two  bracts. 

Bibracteolate  (-atus).  With  two  bract- 
lets. 

Bicallose  (-osus).     With  two  callosities. 

Bicdrinate  (-atus).    Two-keeled. 

Biceps.      |  With  two  supports  or  stalks 

Bicipital.  >  or  two-headed. 

Bicolor.    Two-colored. 

Biconjugate  (-atus).    Twice  paired. 

Bicornis.     Two-horned. 

Bicomute.    Same  as  preceding. 


GLOSSAEY. 


399 


Bicruris.  Two-legged,  or  with  two  sup- 
ports. 

Bidentate  (-atus).  Having  two  teeth. 
(Not  doubly  dentate.) 

Bvluus.     Lasting  two  days  only. 

Biennial  (Biennis).  Of  two  years'  dura- 
tion ;  31. 

Bifarious  (-ius).  Two-ranked ;  in  two 
vertical  rows. 

Biferus.  Double-bearing ;  fruiting  twice 
a  year. 

Bifid  (-idus).  Two-cleft,  to  the  middle 
or  thereabout. 

Biflorous  (-us).     Two-flowered. 

Bifoliate.    Two-leaved. 

Bifdliolute.     Of  two  leaflets. 

Biforate  (-atus).  Having  two  open- 
ings. 

Biformis.    Two-formed ;  in  two  shapes. 

Bifrons.     With  two  faces  or  aspects. 

Bifurcate  (-atus).  Two-forked;  i.e.  of 
two  prongs  or  forks.  But  it  may 
mean  bis  furcatus;  i.e.,  forked  and 
again  forked. 

Bigeminate  (-atus).  Twice  twin  ;  same 
as  Biconjugate. 

Bigener.  The  offspring  of  a  cross  be- 
tween two  genetically  different  plants. 

Bijugate  (Bijugus).  Two-paired,  as  a 
pinnate  leaf  of  two  juga  or  pairs  of 
leaflets. 

Bilabiate  (-atus).     Two-lipped;  247. 

Bildmellate  (-atus),  or  Bilamettar.  Of 
two  plates  or  lamellae. 

Bilobed  (Bilobus),  or  Bilobate.  Of  two 
lobes,  or  cleft  into  two  segments. 

Bilocellate.  Divided  into  two  locelli; 
263. 

Bilocular  (-am).    Two-celled. 

Bimestris.     Lasting  two  months. 

Bimus.  Lasting  two  years  ;  two  years 
old. 

Binary  (-arius).  Consisting  of  two 
members ;  176. 

Binate  (-atus).     In  pairs  or  twos. 

Bini.     Twin,  or  two  together. 

Binodal  (Binodis).     Having  two  nodes. 

Binomial  Nomenclature,  346. 

Biology.  The  natural  history  of  plants 
and  animals,  i.  e.  of  living  things  ;  1. 

Bipalmate  (-atus).  Twice  palmately  com- 
pound. 

Biparous.  Bearing  two;  as  a  cyme  of 
two  rays  or  axes ;  152,  155. 

Bipdrtible  (-ibilis).  Capable  of  division 
into  two  similar  parts. 

Bipartite  (-rtws).  Divided  almost  into 
two  pieces ;  two-parted. 

Bipet.    Same  as  Bicrurit. 


Bipinnate  (-atus).  Doubly  or  twice 
pinnate;  103. 

Bipinndtifid  (-idus).  Twice  or  doubly 
pinnatifid;  100. 

Bipinndtisect  (-MS).  Twice  pinnately  di- 
vided. 

Biplicate  (-atus).  Twice  folded  or  plaited. 

Biporose  (-osus).    Opening  by  two  pores. 

Biradiate  (-atus).    Of  two  rays. 

Birimose  (-osus).    Opening  by  two  slits. 

Bisected  (-us).  Completely  divided  into 
two  parts ;  99. 

Biseptate  (-atus).    With  two  partitions. 

Biserial  (-ialis),  or  Biseriate  (-iatus).  In 
two  series,  one  above  the  other. 

Biserrate  (-atus).  When  serratures  are 
again  serrate ;  doubly  serrate. 

Bisexual.  Haying  both  stamens  and 
pistil ;  hermaphrodite ;  191. 

Bisulcate  (-atus,  Bisulcus).  Two-grooved; 
having  two  furrows. 

Biternate  (-atus).    Twice  ternate. 

Bladdery.    Thin  and  inflated. 

Blade.  The  lamina,  limb,  or  expanded 
portion  of  a  leaf,  &c. ;  85,  245. 

Blastema.  The  budding  or  sprouting 
part  or  point.  First  used  for  the  axis 
of  an  embryo ;  now  used  for  the  ini- 
tial growth  out  of  which  any  organ 
or  part  of  an  organ  is  developed. 

Bloom.  Besides  its  use  as  equivalent  to 
blossom,  it  denotes  the  white  powdery 
and  glaucous  covering  of  the  surface 
of  many  fruits  and  leaves,  of  a  waxy 
nature. 

Boat-shaped.  Of  the  shape  of  a  boat,  of 
the  deeper  sort,  with  or  without  a  keel. 

Bostrychoidal.  Having  the  form  or  char- 
acter of  a  ringlet,  or  Bostryx ;  157. 

Bostryx.  An  uniparous  helicoid  cyme ; 
156. 

Bothrenchyma.  Tissue  of  plants  com- 
posed of  dotted  or  pitted  ducts. 

Botry-cymose.  Racemes  or  any  botryose 
clusters  cymosely  aggregated  ;  159. 

Bdtryose  (-osus),  Botryoidal.  Of  the  ra- 
cemose type;  144,  145,  146,  153. 

Botrys.   The  equivalent  of  Raceme ;  146. 

Botuliformis.     Sausage-shaped. 
Brdchiate    (-idtus).       With    spreading 
arms,  as  branches  (especially  opposite 
and  decussate)  widely  diverging. 
Brachys.     Greek  for  short,  and  used  in 
compounds;    as,  Brachypodus,  short- 
stalked. 

Bract,  Bractea.     The  leaves  (more  of 
less  modified)  of  a  flower-cluster;  118, 
141. 
Sracteate  (-eatus).    Having  bracts. 


400 


GLOSSARY. 


Bracteola,  Bracteole.    See  Bractlet. 

Bracteolate(-atus).  Having  bractlets. 

Bractlet.  A  bract  of  the  ultimate  grade, 
as  one  inserted  on  a  pedicel  or  ultimate 
flower-stalk,  instead  of  subtending  it ; 
141, 142, 160. 

Bracteose  (-osus).  Full  of,  or  with  con- 
spicuous bracts. 

Branches.  Secondary  axes,  or  divi- 
sions of  an  axis;  47. 

Branchlets.  Ultimate  branches  or  divi- 
sions of  an  axis ;  47. 

Breathing-pores.     See  Stomata,  89. 

Bristle.  A  stiff  hair,  or  any  slender 
body  or  outgrowth  which  may  be 
likened  to  a  hog's  bristle. 

Bristly.     Beset  with  bristles. 

Brunneus.     Deep  brown. 

Brush-shaped.     See  Aspergilliform. 

Bryology.    The  botany  of  Mosses. 

Bud.  The  undeveloped  state  of  a  stem 
or  branch,  with  or  without  leaves; 
6,40. 

Bud-scales.   The  teguments  of  a  bud ;  40. 

Bulb  (Bulbus).  A  leaf-bud  (commonly 
subterranean)  with  fleshy  scales  or 
coats ;  43,  62. 

Buibiceps.    A  stem  with  bulbous  base. 

Bulbiferous  (-us).    Bulb-bearing. 

Bulbillus,  Bulbulus.  Diminutive  bulb. 
Same  as 

Jfalblet.  A  small  bulb,  especially  such  as 
is  produced  in  the  air,  in  the  axil  of  or- 
dinary leaves,  or  upon  them ;  63. 

Bulbodium.  A  synonym  of  Corm,  the 
"solid  bulb." 

Bulbo-tuber.     Synonym  of  Corm. 

Bulbous,  Bulbosus.  Having  bulbs  or  the 
structure  of  a  bulb. 

Bullate  (-atus).  Said  of  a  puckered  sur- 
face (as  if  blistered),  thrown  into  por- 
tions which  are  convex  and  projecting 
on  one  side  and  concave  on  the  other. 
Also  used  in  specific  names,  in  its 
more  literal  sense  for'inflated. 

Bursicula.  A  small  pouch  (bursa); 
such  as  that  which  encloses  the  disk 
or  gland  of  the  caudicle  of  the  pollin- 
ium  of  an  Orchis. 

Bursiculatus.  Furnished  with  a  bursi- 
cula  or  pouch. 

Byssaceous  (-eus).  Composed  of  fine 
threads,  like  byssus  or  fine  flax. 


Caducous  (-us).  Dropping  off  very  early, 
as  the  calyx  of  a  Poppy  at  the  time 
of  expansion;  243. 

Carukut.    Sky  blue,  or  pure  blue. 


ius.  Lavender-color ;  pale  green  with 
whitish  or  gray. 

Calathidium,  Cdlathis.  Literally  a  bas- 
ket; a  name  for  the  head  of  flowers  (or 
better  for  the  involucre  only)  of  Com- 
positae. 

Calathiform  (-ormis).  Cup-shaped  ;  of 
somewhat  hemispherical  outline. 

Calcar.  A  spur;  mostly  used  for  the  nec- 
tariferous one  of  a  calyx  or  corolla. 

Cdlcarate  (-dtus).  Furnished  or  pro- 
duced into  a  spur. 

Cdlceolate  (-atus),  or  Calceiformis. 
Shaped  like  a  slipper  or  shoe. 

Callose  (-osus).  Bearing  callosities 
(calli),  or  hard  protuberances. 

Calvus.  Bald,  as  an  akene  without 
pappus. 

Calycdnthemy.  Name  of  the  monstros- 
ity in  which  the  calyx  imitates  an 
exterior  corolla ;  174. 

Calyciflorous  (Calyciflorce),  340. 

Calycine  ( Calycinus).    Relating  to  calyx. 

Calyculate  (-atus).  Bearing  bracts  next 
the  calyx  which  imitate  an  external 
or  accessory  calyx. 

Calyculus.  An  involucre  or  involucel 
imitating  an  additional  calyx. 

Calyptra.  The  hood  or  veil  of  the 
spore-case  of  a  Moss ;  or  some  cover- 
ing body  like  it. 

Calyptrate  (-atus).  Furnished  with  a 
calyptra,  or  something  like  it. 

Calyptriform  (-ormis).  Calyptra-shaped ; 
as  the  calyx  of  Eschscholtzia. 

Calyx.  The  flower-cup,  the  exterior 
perianth  ;  164. 

Cdmara  and  its  diminutive  Cnmerula 
(chamber)  are  sometimes  used  for  the 
cells  of  a  fruit. 

Cambium.  Old  name  of  the  viscid  mat- 
ter between  bark  and  wood  in  com- 
mon trees  or  shrubs  in  spring;  now 
used  for  the  nascent  structure  there 
forming,  or  Cambium  layer ;  78. 

Campdnulate (-atus).  Bell-shaped;  elon- 
gated cup-shaped  or  shorter,  and  broad 
from  the  base ;  249. 

Campaniformis.    Same  as  Campanulate. 

Campylospermous  (-^us).  Curved-seeded. 
Said  of  seed-like  fruits  or  carpels,  as 
those  of  some  Umbelliferas,  in  which 
the  contained  seed  is  involute  by  the 
lateral  edges,  so  as  to  produce  a  longi- 
tudinal furrow  on  the  ventral  face. 

Campylotropous  (-us),  or  less  correctly 
Campy  lotropal,  or  Campulitropous. 
An  ovule  or  seed  which  is  curved 
in  its  formation  so  as  to  bring  the 


GLOSSARY. 


401 


micropyle  or  true  apex  down  near  to 
the  hilum;  279. 
Canalicaulate    (-atus).     Channelled,  or 

with  a  longitudinal  groove. 
Cdncellate  (-dtus).   Latticed;  resembling 

lattice-work. 
Cdndidus.     Pure  white. 
Canescens.      Hoary,   usually  with  gray 

pubescence. 

Canus.     Gray-white;   whiter  than  the 
preceding. 

\  So  slender  that  it 
Capillaceous(-eus).  I  may  be  compared 
Capillary  (-am).  f  with  the  hairs  of 

)  animals. 

Capitate  (-atus).     Head-shaped,  or  col- 
lected in  a  head;  147. 
Capitellate  (-atus).     Diminutive  of  Cap- 
itate. 

Capitulum.    A  head  of  or  simple  globu- 
lar cluster  of  sessile  flowers;  147. 
Capreolate   (-atus).     Bearing  a   tendril 

(capreolus). 

Capsule  (-ula).     A  dry  and  dehiscent 
pericarp  composed  of  more  than  one 
carpel ;  289,  293. 
Capsular.     Of  the  nature  of,  or  relating 

to,  a  capsule. 

Capsuliferous.     Capsule-bearing. 
Carcerulus.    An  unused  name  for  an 
indehiscent    and    several-celled    dry 
fruit;  297. 

Carina.    A  keel;   used  either  for  the 
two  combined  lower  petals  of  a  papil- 
ionaceous corolla  (185);   or  for  a  sa- 
lient longitudinal   projection   on   the 
centre  of  the  lower  face  of  an  organ, 
as  on  the  glumes  of  many  Grasses. 
Cdrinate  (-atus).     Keeled. 
Cariopsis  or  Caryopsis.   A  grain ;  a  seed- 
like  fruit  with  thin  pericarp  adnate  to 
the  contained  seed ;  295. 
Carneus.     Flesh-colored,  very  pale  red. 
Caro.    Flesh,  as  the  pulp  of  a  melon,  or 

the  fleshy  part  of  a  drupe. 
Carpadilium.     Synonym  of  Cremocarp. 
Carpel,  Carprtlum.    A  simple  pistil,  or 
an  element  of  a  compound  pistil,  an- 
swering to  one  leaf ;  167.  260. 
Carpid,  Carpidium.    Synonym  of  carpel. 
Carpolngy.     The  botany  of  fruits. 
Carpophore  (CarpSphorum).     A  portion 
of  receptacle   prolonged   between  the 
carpels;  212. 
Cdrpophyll  (Cnrpoplyllum).      Literally 

fruit-leaf;  synonym  of  Carpel ;  260. 
Cartilaginous  or    Cartilayineous  (-em). 
Of  the  texture  of  cartilage  or  gristle ; 
firm  and  tough. 


Caruncle  (Canimcula).  An  excrescence 
at  or  about  the  hilum  of  certain  seeds ; 
308. 

Caryophyllaceous(-eus).  Resembling  or 
relating  to  the  corolla  of  Dianthus 
Caryophyllus  (246),  or  to  the  Pink 
family. 

Caryopsis.     See  Cariopsis. 
Cassideus.    Helmet-shaped. 
Cassus.    Empty,  as  an  anther  contain- 
ing no  pollen. 
Castrate    (-atus).     Said    of    a    stamen 

which  wants  the  anther. 
Catapetalous  (-M«).     Where   petals   are 
united  only  by  cohesion  with  united 
stamens,  as  in  Mallow. 
Cataphylla.      Answers  to  the   German 
"  Niederblatter,"    or     under-leaves, 
those  at  the  beginning  of  a  growth, 
cotyledons,  bud-scales,  scales  on  rhi- 
zomes, &c. ;  6. 

Catenulate  (-dtus).  Formed  of  parts 
united  end  to  end,  like  the  links  of  a 
chain. 

Catkin.   A  scaly  spike  (see  Ament);  150. 
Caudate  (-atus).     Furnished  with  a  tail 
(cauda),  or  with  a  slender  tip  or  ap- 
pendage resembling  a  tail. 
Caudex.    A  trunk  or  stock  of  a  plant ;  50. 
Caudide  (Caudicula).     The  stalk  of  a 

pollinium,  &c. 
Caulescent  (-ens).    Having  an  obvious 

stem. 

Caulicle  (Cauliculus).  The  initial  stem 
in  an  embryo,  generally  named  the 
Radicle;  10. 

Cauline  (-inus).    Belonging  to  the  stem. 
Caulis.    Greek  form  Caulon.    The  stem 

of  a  plant. 

Caulocdi-pic  or  Caulocarpous.     Applied 
to  plants  which  live   to  flower  and 
fructify  more    than   once  or  indefi- 
nitely. 
Caulome,  Cauldma.     The  stem-part  of  a 

plant. 

Cephalanthium.  One  of  the  names  of 
the  head  or  capitulum  in  Composite ; 
148. 

Cell  ( Cellula).  The  anatomical  element 
of  plants;  28.  The  cavity  of  an 
anther  which  contains  the  pollen, 
or  an  anther-lobe,  thus  taken  in  the 
sense  of  the  circumscribing  wall  as 
well  as  the  cavity;  251,  254.  The 
cavity,  or  any  one  cavity  of  an  ovary 
or  pericarp,  containing  the  ovules  or 
seeds;  262. 

Cellular  Plants,  Cellulares,  340. 
Cellide  (-wio).    Diminutive  of  cell;  of 

LIBRARY 

STATE  TEACHERS  COLLFO, 
SANTA  BARBARA.  CALIFO*N'/ 


402 


GLOSSARY. 


the  same  meaning  as  Cell  in  vegetable 
anatomy;  28. 

Cellulose.  The  material,  chemically  con- 
sidered, of  which  the  wall  of  the  cell 
consists. 

Cenobium.  A  name  of  the  peculiar  four- 
parted  fruit  (or  the  four  nutlets  around 
a  common  style)  which  distinguishes 
Labiatse  and  Borraginacese. 

Centrifugal.  Tending  or  developing 
from  the  centre  outward. 

Centripetal.  Tending  or  developing  from 
without  toward  the  centre. 

Cephalanthium.  Synonym  of  Antho- 
dium. 

Ceratium.  A  siliquifonn  capsule,  such 
as  that  of  Corydalis,  Cleome,  &c. 

Cereal.  Belonging  to  corn  and  the 
allied  grains. 

Cerinus.    Of  the  color  of  wax. 

Cemuous  (-MM*).    Nodding. 

Chceta.    Greek  for  a  bristle,  Latin  Seta. 

Chaff".  Small  scales;  dry  and  depau- 
perate bracts ;  such  as  those  on  the  re- 
ceptacle of  a  sunflower  and  many  other 
Compositae  ;  also  glumes  of  Grasses. 

Chaffy.  Provided  with  or  having  the 
texture  of  chaff. 

Chaldza.  The  part  of  an  ovule  where 
coats  and  nucleus  are  confluent;  277. 

Channelled.  Hollowed  out  longitudi- 
nally like  a  gutter.  See  Canaliculate. 

Character.  A  diagnostic  description, 
or  the  enumeration  of  essential  differ- 
ences; 361. 

Chasmdgamy.  The  opening  of  the  peri- 
anth at  flowering  time;  the  opposite 
of  Cleistogamy. 

Chartaceous  (-eus).  Having  the  texture 
of  writing-paper. 

Chlorophyll.  The  green  matter  of  leaves 
and  other  vegetation  ;  76,  88. 

Chloros.  Greek  for  green.  Enters  into 
compounds,  such  as  Chloranthiis, 
green-flowered,  Chloranthy,  same  as 
Chlorosis,  as  when  petals  turn  green; 
172. 

Chlorosis.  Literally  becoming  green,  as 
some  flowers  in  retrograde  metamor- 
phosis. Also  used  contrariwise  for 
the  loss  of  a  normal  green  color ;  172. 

Chorda  Pistlllaris.  A  line  of  tissue  reach- 
ing from  stigma  to  ovary. 

Choripetalous  (-us).  Same  as  Polypeta^ 
bus,  i.  e.  petals  unconnected  ;  244. 

Chorisepalous.  Same  as  Polysepalotis,&c. 

Chorisis.  The  separation  of  a  leaf  or 
phyllum  into  more  than  one  ;  202. 

Chorittophyllut.    Separate-leaved. 


Chromule  (^ula).  Coloring  matter  of 
plants  other  than  chlorophyll,  espe- 
cially that  of  petals. 

Chrysos.  Greek  for  golden,  or  golder- 
yellow;  as 

Chrysanthus.     Yellow-flowered,  &c. 

Cicatrix,  Cicatricula.  A  scar  left  by 
the  fall  of  a  leaf  or  other  organ. 

Ciliate  (-atus),  Ciliaris.  Marginally 
fringed  with  hairs. 

Cilium,  pi.  cilia.  Marginal  hairs,  form- 
ing a  fringe,  like  the  eyeJash.  (The 
name  has  been  extended  in  scientific 
books  to  undividual  hairs,  and  of  a 
surface  as  well  as  edge.) 

Cincinnus.  A  curl :  name  of  a  uniparous 
scorpioid  cyme,  which  is  Cincinnal; 
156,  157. 

Cinenchyma.    Laticiferous  tissue. 

Cinerascens,  Cintraceus.    Ash-grayish. 

Cinereous  (-eus).     Ash-gray. 

Cinnabarinus.  Cinnabar-color  ;  scarlet 
touched  with  orange. 

Circinal  (-alis).  Involute  from  the  tip 
into  a  coil;  133. 

Circinate,  or  Circinnate  (-atus).  Same 
as  preceding;  or  sometimes  meaning 
coiled  into  a  ring  only. 

Circumscissile,  or  Circumcissile  ( Circum~ 
scissus).  Cut  circulary  and  transverse- 
ly ;  divided  transversely ;  291,  293. 

Circumscription  (-io).  The  general  out- 
line of  the  margin  of  a  flat  body. 

Cirrhiferous  (-us)  and  Cirrhose.  Ten- 
dril-bearing. 

Cirrhus.    A  tendril ;  54. 

Citreus,  Citrinus.    Lemon-colored. 

dados.  Greek  for  branch ;  whence  such 
terms  as 

Cladodium.     Same  as  Cladophyllum. 

Clddophyll,  Cladophylla.  Branches  as- 
suming the  form  and  function  of  foli- 
age ;  65,  66. 

Clavate  (-atus),  Claviformis.  Club- 
shaped. 

Clavellate.     Diminutive  of  Clavate. 

Claviculate  (-atus).  Furnished  with  clav- 
icuke;  viz.,  tendrils,  hooks,  or  other 
appliances  for  climbing. 

Class,  325. 

Classification,  315. 

Clathrate  (-atus).     Latticed. 

Claw.  The  narrowed  base  or  stalk  which 
some  petals,  &c.,  possess;  245. 

Cleistdyamy,  Cleistogamous,  Cleistogamic. 
Close-fertilization  in  unopened  blos- 
soms ;  241. 

Cleistogeny,  Cleistogfnous,  241.  Same  as 
Cleistogamy. 


GLOSSARY. 


403 


Cleft.  Cut  half-way  down  or  there- 
about; 98. 

Climbing.  Rising  by  laying  hold  of 
surrounding  objects  for  support;  51. 

Clinandrium.  The  anther-bed  in  Orchi- 
daceae. 

Clinanthium.  A  name  for  the  receptacle 
of  inflorescence  in  Composite ;  148. 

Clinium.  Used  in  Greek  compounds  for 
receptacle,  e.g.  Periclinium,  for  an 
involucre  around  the  receptacle  of 
inflorescence. 

Close-fertilization.  Fecundation  by  own 
pollen ;  216,  280. 

Cloves.  A  gardener's  name  for  young 
bulbs  developed  around  a  mother  bulb. 

Club-shaped.  Gradually  thickened  up- 
ward from  a  slender  base. 

Clustered.  Collected  in  a  bunch  of  any 
sort.  Cluster  is  a  good  indefinite  name 
for  any  assemblage  of  flowers  on  a 
plant. 

Clypeate  (-atus),  Clypeiformis.  Buck- 
ler-shaped. 

Coacerrate  (-atus).    Heaped  together. 

Coddnate  (-atus),  Coadunatus.  Same  as 
Adnate. 

Coalescence.  Union  of  similar  parts; 
179,  180. 

Coalescent  (-ens),  Coalitus.  Cohering; 
properly  applied  to  the  organic  cohe- 
sion of  similar  parts. 

Coarctate  (-atus).    Crowded  together. 

Coated.  Composed  of  layers  as  an 
onion,  or  furnished  with  a  covering 
or  rind. 

Cobwebby.   Bearing  long  and  soft  entan- 


Coccineus.  Bright  red  or  scarlet  (red 
with  a  little  yellow). 

Coccus.  Greek  for  a  kernel  or  nutlet, 
from  which  the  Latin  Coccum,  the 
kermes  or  scarlet  grain  (supposed 
berry)  of  the  Quercus  coccifera;  used 
botanically,  mostly  in  the  form  of 
"coccus,"  for  the  portions  into  which 
a  schizocarp,  or  lobed  fruit  with  one- 
seeded  cells,  splits  up:  these  portions 
are  Cocci  or  Coccutes;  296. 

Cochlear  (Cochlearis).  Spoon-shaped. 
Unmeaning!}'  applied  also  to  a  form 
of  imbricative  aestivation  with  one 
piece  exterior;  137. 

Cochleate  (-atus).  Shell-shaped,  i.  e. 
spiral  in  the  manner  of  a  snail-shell. 

Ccelospermous  (-tis).  Hollow-seeded;  ap- 
plied to  seed-like  carpels  of  Umbelli- 
ferae  with  ventral  face  incurved  at  top 
and  bottom,  as  in  Coriander. 


Cananthium.   Synonym  of  Clinanthium. 

Ccenobio.    Synonym  of  Carcerulus. 

Cohesion.  The  congenital  union  of  one 
organ  with  another  ;  either  similar 
parts  (coalescence),  or  dissimilar  parts 
(adnation). 

Cohort.  In  classification  a  group  next 
superior  to  order,  326. 

Coleorhiza.  Root-sheath;  the  invest- 
ment (belonging  to  the  cotyledon  orj 
plumule)  through  which  the  primary' 
root  in  many  Monocotyledons  bursts 
in  germination ;  26. 

Collar  ( Collum).  Name  of  an  imaginary 
something  intermediate  between  pri- 
mary stem  and  root. 

Collateral.    Standing  side  by  side. 

Collective  Fruits.  The  aggregation  of 
the  fruits  of  several  flowers  into  one 
mass;  301. 

Colored.  Of  other  color  than  the  green 
of  herbage;  118. 

Columella.  The  persistent  axis  of  cer- 
tain capsules,  spore-cases,  &c. ;  289. 

Column  (Columna).  Body  formed  by 
the  union  of  the  filaments  among  them- 
selves (as  in  a  malvaceous  flower),  or 
with  the  style  or  stigma,  as  in  Or- 
chids; 250. 

Columnar.  Column-shaped  ;  pillar* 
shaped. 

Coma.  Literally  a  head  of  hair;  a  tuft 
of  hairs  of  any  sort ;  specially  a  toft  of 
hairs  on  a  seed ;  306.  Also  the  name 
of  the  whole  head  of  a  tree. 

Commissure  (-ura).  The  face  by  which 
two  carpels  cohere,  as  in  Umbelliferae. 

Common  ( Communis).  General  or  prin- 
cipal, as  opposed  to  partial. 

Comose  (-osus),  sometimes  Comatus. 
Furnished  with  a  coma. 

Ctmplanate  (-atus).    Flattened. 

Complete  (Completus).  Having  all  the 
parts  belong  to  it  or  to  the  type ;  175.  i 

Complicate  (-atus).    Folded  upon  itself. 

Compound.  Said  of  similar  parts  aggre- 
gated into  a  common  whole.  Com- 
pound Flower,  147.  Compound  Pistil, 
263.  Compound  Inflorescence,  159. 

Compound  Leaf.  One  divided  into  sep- 
arate blades;  100. 

Compressed  (-us).   Flattened  lengthwise. 

Concaulescence.  A  name  for  the  coales- 
cence of  axes ;  158. 

Conceptacle  (-aculum).  Originally  used 
by  Linnaeus  for  what  is  now  called 
Follicle ;  and  later  for  the  pair  of  fol- 
licles of  Asclepiadaceae  and  Apocy- 
nacese: 


404 


GLOSSABY. 


Conchiformis.  Shaped  like  one  valve  of 
a  bivalve  shell. 

Concinnus.    Neat  or  elegant. 

Concolor.  Of  the  same  or  of  uniform 
color. 

Conduplicate  (-atus),  Conduplicativus. 
Folded  together  lengthwise;  133. 

Cone.    See  Strobile. 

Confertus.    Closely  packed  or  crowded. 

Conferruminate  (-atus).  Stuck  together 
by  adjacent  faces,  as  the  cotyledons  of 
Horsechestnut ;  314. 

Confluent  (-ens).  Blended  into  one;  pass- 
ing by  degrees  the  one  into  the  other. 

Conformed  (-ormis).  Similar  to  in  form ; 
or  closely  fitted  to,  as  a  seed-coat  to 
the  nucleus. 

Congested  (-ws).    Crowded  together. 

Conglobate  (-atus).    Collected  into  a  ball. 

Conglomerate  (-atus).  Densely  clus- 
tered or  heaped  together. 

Coniferous  (-M»).    Cone-bearing. 

Conjugate  (-atus).  Coupled;  in  single 
pairs.  Conjugate-pinnate,  104. 

Connate  (-atus).  United  congenitally ; 
107,  182. 

Connate-perfoliate.  United  at  base  in 
pairs  around  the  supporting  axis ;  108. 

Connective  (-ivum).  A  portion  of  a  sta- 
men which  connects  the  two  cells  or 
lobes  of  an  anther;  251. 

Connivent  (-ens).  Coming  into  contact 
or  converging. 

Conocarpium.  An  unused  name  for  an 
aggregate  fruit,  such  as  a  strawberry, 
consisting  of  many  carpels  on  a  coni- 
cal receptacle ;  298. 

Consolidated.  When  unlike  parts  arc 
coherent. 

Continuous.  The  reverse  of  articulated 
or  interrupted. 

Contorted  (-us).  Twisted;  or  bent  or 
twisted  on  itself.  In  ^Estivation,  the 
same  as  Convolute ;  138. 

Contortuplicate  (-atus).  Twisted  and 
plaited  or  folded. 

Contracted.  Either  narrowed  or  short- 
ened. 

Contrary  (-arius).  Opposite  in  direction 
to  the  part  compared  with ;  as  a  silicle 
compressed  contrary  to  the  dissepi- 
ment. 

Convolute  (-utus)  or  Convolutive  (-ivus). 
Rolled  up  from  the  sides  or  longitudi- 
nally. In  ^Estivation,  138.  In  Ver- 
nation, 133. 

Ciralloid  (-eus).    Coral-like. 

C6rculum.  Old  name  for  the  embryo, 
or  Corteminis;  311. 


Cordate  (-atus),  sometimes  Cordiform 
(-ormis).  Heart-shaped ;  like  the  fig- 
ure of  a  heart  on  cards ;  the  stalk  at 
the  broader  and  notched  end  ;  96. 

Coriaceous  (-eus).  Leathery  in  consist- 
ence. 

Cork,  81. 

Corky.    Of  the  texture  of  cork. 

Corky  Envelope,  76. 

Corm  (Cormus).  A  bulb-like  fleshy 
stem,  or  base  of  a  stem  ;  a  "  solid 
bulb ;  "  61. 

Cormophytes  ( Cormophyta),  341. 

Corneous  (-eus).    Of  the  texture  of  horn. 

Corniculate  (-atus).  Furnished  with  a 
little  horn. 

Cornu.  A  horn ;  i.  e.  a  horn-like  process ; 
sometimes  used  for  Calcar,  a  spur. 

Cornute(-utus).  Furnished  with  a  horn- 
like process  or  spur. 

Cordtta.  The  interior  perianth,  com- 
posed of  petals;  16»,  243. 

Corollaceous  (-eus),  Carollinus.  Pertain- 
ing to,  or  resembling  corolla. 

Corolliferous  (-us).     Bearing  a  corolla. 

Corolliflorous,  Corolliflorce,  340. 

Corollula.     Diminutive  of  corolla. 

Cordna.  A  crown ;  an  inner  appendage 
to  a  petal,  or  to  the  throat  of  a  corolla ; 
210,  246.  Or  any  coronet-like  append- 
age at  the  summit  of  (crowning)  an 
organ. 

C6ronate  (-atus).  Crowned,  having  a 
corona,  &c. 

Cordniform  (-ormis).  Shaped  like  a 
crown  or  coronet. 

Corrugate  (-atus  or  -ativus).  Wrinkled 
or  in  folds ;  133. 

Cortex.     Rind  or  bark. 

Cortical  (-alis).     Relating  to  bark. 

Corticate  (-atus).  Coated  with  a  bark 
or  with  an  accessory  bark-like  cover- 
ing. 

C6rymb  (Corymbus).  A  flat-topped  or 
merely  convex  and  open  flower-cluster 
of  the  indeterminate  or  centripetal 
order;  146. 

Corymbiferous  (-us).    Bearing  corymbs. 

Corymbose.  In  corymbs,  or  in  the  man- 
ner of  a  corymb.  The  corymb  of 
Linnaeus  and  of  other  writers  down 
to  Roeper  included  most  cymes.  So 
that  much  cymose  inflorescence  is 
in  descriptions  loosely  said  to  be 
corymbose,  or  a  stem  is  said  to  be 
corymbosely  branched,  even  when 
the  evolution  is  centrifugal ;  146. 

Costa.  A  rib;  when  single,  a  midrib 
or  mid-nerve. 


GLOSSARY. 


405 


Costal-nerved.  With  nerves  springing 
from  a  midrib ;  92. 

Costate  (-atus).  Ribbed;  furnished  with 
one  or  more  longitudinal  primary 
veins  or  ribs. 

Cotyledons  (  Cotyledon,  pi.  Cotyledones). 
The  "  seed-lobes,"  being  the  leaves  or 
first  leaves  of  the  embryo;  viz.,  the 
one,  or  the  pair,  or  rarely  the  whorl  of 
leaves  borne  by  the  radicle  or  caulicle ; 
10,  311,  313. 

Cotyliformis.  Dish-shaped,  or  wheel- 
shaped  with  an  erect  or  ascending 
border. 

Crateriform  (-ormis).  In  the  shape  of 
a  goblet  or  cup,  of  hemispherical  con- 
tour or  more  shallow ;  248. 

Crdmocarp  ( Cremocdrpium).  A  dry  and 
seed-like  fruit,  composed  of  two  one- 
seeded  carpels,  invested  by  an  epigy- 
nous  calyx,  and  separating  at  matu- 
rity; 297. 

Creeping.  Running  along  or  under 
ground  and  rooting;  53. 

Crena,  Crenatura.  A  rounded  tooth  or 
notch. 

Crenate  (-atus).  Toothed  by  crenatures ; 
scalloped;  98. 

Crenel,  Crenelled.  Same  as  Crenature 
and  Crenate. 

Crenulate  (-atus).  Diminutive  of  Cre- 
nate, t.  e.  with  small  crenatures. 

Crested.  Furnished  with  any  elevated 
line,  ridge,  or  conspicuous  elevation 
on  the  surface,  especially  such  as  may 
be  likened  to  the  crest  of  a  helmet. 

Cretaceus.    Chalk-white;  chalky. 

Cribrose  (-osus)  and  Cribriform  (-ormis). 
Pierced  like  a  sieve. 

Cribriform  Cells,  77. 

Crinitus.  Bearded  with  long  and  weak 
hairs. 

Crispatus.    Curled  or  crispy. 

Cristate  (-atus).    Crested. 

Croceus,  Crocatus.  Saffron-colored,  i.  e. 
deep  reddish-yellow. 

Cross-breeds.  The  progeny  of  interbred 
varieties ;  321. 

Cross-fertilization.  Fecundation  by  pol- 
len of  another  flower  and  of  another 
individual;  216. 

Crown.    See  Corona,  210;  246. 

Crowned.    See  Coronate. 

Crowning  ( Coronans).  Borne  on  the 
summit  of  an  organ. 

Cruciate  (-atus),  Cruciform  (-ormis). 
Cross-shaped. 

Cruciferous  (-us).  Cross-bearing;  used 
in  the  sense  of  Cruciform  ;  as  the 


"cruciferous"    corolla  of  the  order 

Cruciferse;  246. 
Crumpled.    See  Corrugate. 
Crustaceous  (-us).     Of  hard  and  brittle 

texture. 

Cryptos.    Greek  for  concealed ;  whence 
Cryptogamia.     Cryptogamous  or  Cryp- 

togamic  plants  ;  3,  335,  344. 
Cryptogamous.    Pertaining  to  the  above. 
Cucullate    (-atus),     Cuculldris,    Cuculli- 
formis.      Hooded,     or    hood-shaped, 

cowled. 
Culm  (  Culmus).    The  peculiar  stem  or 

straw  of  Grain-plants  and  Grasses ;  50. 
Cultrate  (-atus),  Cultriformis.     Shaped 

like  a  broad  knife-blade. 
Cuneate  ( Cuneatus),  Cuneiform  (-ormis). 

Wedge-shaped ;    triangular  with    an 

acute  angle  downward ;  95. 
Cup-shaped.    In  the  form  of  a  drinking- 

cup. 
Cupule  ( Cupula).    The  acorn-cup    and 

the  like;  296. 
Cupularis,  Cupulatus.     Furnished  with 

or  subtended  by  a  cupule  or  any  re- 
sembling body. 

Cupuliferous  (-MS).     Cupule-bearing. 
Curvinerved  (-ius).    When  the  ribs  of  a 

leaf  are  curved  in  their  course ;  92. 
Curviserial.   In  curved  or  oblique  ranks ; 

124. 

Cushion.    The    enlargement  at  or  be- 
neath the  insertion  of  many  leaves. 
Cuspidate  (-atus).    Tipped  with  a  Cusp, 

or  sharp  and  rigid  point ;  97. 
Cut.    Same  as  incised,  or  in  a  general 

sense  as  cleft. 
Cuticle  (  Cuticula).    The  outermost  skin 

or  pellicle. 
Cutting.    A  severed  portion  of  a  plant 

used  for  bud-propagation ;  43. 
Cydneus.    A  clear  bright  blue. 
Cydthiform  (-ormis).  Cup-shaped;  in  the 

form  of  a 
Cyaikus.     A   drinking-cup,  such  as   a 

goblet  or  wine-glass. 
Cycle.    A  circle.     Sometimes  used  for 

one  turn  of  a  helix  or  spire;  122. 
Cyclical.    Relating  to  a  cycle ;  or  coiled 

into  a  circle;  119,  120. 
Cylindraceous.      Somewhat    or    nearly 

cylindrical. 
Cylindrical  (-««).     Elongated  and  with 

circular  cross-section ;  in  the  form  of  a 

cylinder. 
Cymbaform    or     Cymbiform    (-ormis). 

Boat-shaped. 
Cyme  (  Cyma).    A  flower-cluster  of  the 

determinate     or     centrifugal     type. 


406 


GLOSSARY. 


especially  a  broad  and  flattish  one; 
151. 

Cymo-btitryose.  When  cymes  are  ar- 
ranged in  botryose  manner;  159. 

Cymose  (-osus).  Bearing  cymes,  or  re- 
lating to  a  cyme ;  151. 

Cymule  ( Cymula).  Diminutive  cyme, 
or  a  portion  of  a  cyme ;  151. 

Cynarrhddium,  Name  of  such  a  fruit  as 
that  of  the  Rose ;  fleshy,  hollow,  and 
enclosing  achenia. 

Cypsela.  Name  of  an  achenium  in- 
vested by  an  adnate  calyx,  as  the  fruit 
of  Composite;  295. 

Cystolith.  One  of  the  mineral  and 
usually  partly  crystalline  concretions 
of  the  cells  of  the  epidermis  of  or 
subjacent  tissue  of  the  leaf  in  various 
plants,  especially  in  Urticaceae. 

Cytoblast.  An  obsolete  name  for  the 
nucleus  of  a  cell  of  cellular  tissue. 


Ddctylose  (-osus).  Fingered,  or  finger- 
shaped. 

Dasyphyllous  (-us).    Woolly-leaved. 

Dealbate  (-atus).  Whitened  over  (as  if 
whitewashed)  with  a  white  powder  or 
minute  pubescence. 

Deca.  Greek  for  ten,  .compounded  with 
various  words,  such  as 

Decagynia.  One  of  the  Linnsean  artificial 
orders;  337. 

Decdgynous  (-us).  With  ten  styles  or 
carpels. 

Decdmerous  (-MS).  Of  ten  members ;  176. 

Decandria.  A  Linnsean  class  with  ten 
stamens;  334. 

Decandrous  (-us).  With  ten  stamens; 
249. 

Decapetalous  (-us),  Decasepalous,  &c. 
With  ten  petals  or  sepals,  &c. 

Deciduous  (-ws).  Falling,  or  subject  to 
fall  in  season,  as  petals  after  anthesis, 
and  leaves  (except  of  evergreens)  in 
autumn;  243. 

Declinate  (-atus),  or  Declined.  Bent  or 
curved  downward  or  forward. 

Decompound.  Several  times  compound- 
ed or  divided;  102,  104. 

Decompositus.     Decompound. 

Decumbent  (-ens).  Reclining,  but  with 
summit  ascending;  53. 

Decurrent  (-ens),  Decursive.  Running 
down  into ;  as  where  leaves  are  seem- 
ingly prolonged  below  their  insertion, 
and  so  run  down  the  stem. 

Decussate  (-atus).  In  pairs  alternately 
crossing  at  right  angles. 


Deduplication,  Fr.  Dedoublement.  Same 
as  Chorisis ;  202. 

Definite  (-itus).  Of  a  fixed  number,  not 
exceeding  twenty ;  or  of  a  fixed  order. 

Definite  Inflorescence.  Where  axes  of 
inflorescence  end  in  a  flower ;  144, 151. 

Deflexed  (-us).  Bent  or  turned  abruptly 
downward. 

Deflorate  (-atus).  Past  the  flowering 
state. 

Defoliate  (-atus).  Having  cast  its  leaves. 

Defoliation,  87. 

Dehiscence  (-entia).  The  mode  of  open- 
ing of  a  capsule  or  anther  by  valves, 
slits,  or  regular  lines ;  288. 

Dehiscent  (-ens).  Opening  by  regular 
dehiscence;  292. 

Deliquescent  (-ens).  Dissolving  or  melt- 
ing away,  as  a  stem  divided  into 
branches;  48. 

Deltoid  (-oides).  Having  the  shape  of 
the  Greek  letter  A. 

Demersed  (-us).  Under  water ;  same  as 
submersed. 

Dendritic  (-icus),  Dendroid  (-oideus). 
Tree-like. 

Dendron.     Greek  for  tree. 

Deni.    Ten  together. 

Dens.    A  tooth. 

Dentate  (-atus).  Toothed;  specially  with 
salient  teeth  not  turned  forward ;  98. 

Denticulate  (-atus).  Minutely  toothed; 
having  denticulations,  or  diminutive 
teeth. 

Denudate  (-atus).  Made  naked ;  stripped. 

Deorsum.    Downwards. 

Depauperate  (-atus).  Impoverished;  as 
if  starved;  or  diminutive  for  want  of 
favorable  surroundings. 

Depressed  (-us).  Having  the  appear- 
ance or  shape  as  if  flattened  from 
above. 

Derma.  Greek  for  skin  or  surface  of  a 
plant  or  organ. 

Descending  (-ens).  Tending  or  turning 
gradually  downward. 

Descending  Axis.    Primary  root ;  11. 

Determinate.  Limited  in  number  or  ex- 
tent; as  are  the  axes  of  determinate 
inflorescence;  144,  151. 

Desinens.    Terminating  in. 

Desmos.  Greek  for  things  bound,  or  as  if 
chained  together. 

Dextrorse  (Dextrorsus:  adv.  Dextror- 
sum).  Toward  the  right  hand,  or  re- 
lating to  it ;  51,  140. 

Di,  Dis.  In  Greek  compounds,  two,  or 
double. 

Diachenium.    Synonym  of  Cremocarp. 


GLOSSARY. 


407 


Diadelphia.     A    Linntean    (335)  class 

having  the  stamens. 

Diadelphous  (-us).  Combined  by  their 
filaments  into  two  sets ;  250. 

Diagnosis.  A.  brief  distinguishing  char- 
acter. 

Dialypetalas,  341. 

Dialypetalous  (-us).  Same  as  polypeta- 
lous,  f .  e.  of  separate  petals ;  244. 

Dialyphyllous  (-us).  Bearing  separate 
leaves. 

Diandria.  A  Linnaean  class  with  per- 
fect flowers  having  only  two  stamens ; 
334. 

Diandrous  ( Diander,  &c. ).  Having  two 
stamens;  249. 

Diaphanous  (-MS).  Letting  the  light 
shine  through. 

Dicarpellary.  Composed  of  two  carpels 
or  pistil-leaves ;  261. 

Dichdsium.  A  two-parted  or  two-rayed 
cyme;  152,  155. 

Dichlamydeous  (-eus).  Having  a  double 
perianth;  191. 

Dichotomous  (-us).  Forked  in  pairs ; 
two-forked. 

Dichdgamous  (-us),  Dichogamy.  Her- 
maphrodite with  one  sex  earlier  de- 
veloped than  the  other  in  the  blossom ; 
219. 

Diclesium.  Name  of  a  fruit  consisting 
of  an  achenium  within  a  separate  and 
free  covering  made  of  perianth,  as  that 
of  Mirabilis. 

Diclinous  (Didinis).  When  flowers  are 
of  separate  sexes ;  191. 

Dicoccous  (-ws).    Fruits  of  two  cocci. 

Dicotyledons,  Dicotyledones.  Plants  of 
the  class  marked  by  having  two  coty- 
ledons; 27,339,340,344. 

Dicotyledonous  (-eus).  Having  a  pair  of 
cotyledons;  10,  314. 

Didymous  (-us).    Twin,  found  in  pairs. 

Didyndmia.  The  Linnsean  class  marked 
by  didynamy  (335),  i.  e. 

Didynamous  (-us).  When  a  4-androus 
flower  has  the  stamens  in  two  pairs, 
and  one  pair  shorter  than  the  other ; 
250. 

Dieresilis.  Mirbel's  name  for  a  dry 
fruit  composed  of  several  cells  or  car- 
pels connate  around  a  central  axis, 
and  separating  at  maturity,  as  that  of 
Mallow. 

Difformis.    Of  unusual  formation. 

Diffuse  (^usus).  Widely  or  loosely 
spreading. 

Diffamous  (-us).  Of  two  sexes  in  the 
same  cluster. 


Digitate  (-atus).  Fingered ;  a  compound 
leaf  in  which  all  the  leaflets  are  borne 
on  the  apex  of  the  petiole;  101. 

Digitately.  In  a  digitate  mode;  same 
as  Palmately. 

Digitate-Pinnate,  104. 

Digynia.  A  Linnsean  order  character- 
ized by  having  the  gynoscium 

Digynous.  With  two  separate  styles  or 
carpels ;  261. 

Dimerous  (-us)  Of  two  members  in  each 
circle;  176. 

Dimidiate  (-atus).  Halved,  or  as  if  one- 
half  was  wanting. 

Dimorphous  (-us),  Dimorphic,  Dimor- 
phism. Occurring  under  two  forms; 
225,  234. 

Di&cia.  Linnaean  class  (355)  of  plants 
with  the  flowers 

Dioecious  (Dicecius,  Dioicous).  Unisex- 
ual, and  the  two  sexes  borne  by  dis- 
tinct individuals ;  191. 

Diaecio-polygamous.  When  some  indi- 
viduals bear  unisexual  and  others  bi- 
sexual flowers. 

Dipetalous  (-ws).     Two-petaled ;  244. 

Diphyllous  (-us).     Two-leaved ;  243. 

Diplo.     See  Duplo. 

Diplostemonous,  Diplostemony.  Having 
twice  as  many  stamens  as  petals  or 
sepals ;  177,  198. 

Diplotegium.  A  capsule  or  other  dry 
fruit,  invested  with  adnate  calyx ;  an 
inferior  capsule. 

Dipterous  -(us).    Two-winged. 

Diremption  (-io).   Syn.  of  Chorisis;  202. 

Disciferous  (-us).     Disk-bearing. 

Disciform  (-ormis).  Depressed  and  cir- 
cular, like  a  disk  or  quoit. 

Discoidal  or  Discoid  (Discoideus).  Ap- 
pertaining to  a  disk.  A  discoid  head 
is  one  destitute  of  ray-flowers. 

Disc  or  Disk  (Discus).  A  word  used  in 
several  senses.  The  disk  or  dac  of  a 
flower  is  a  development  oT  the  torus 
within  the  calyx,  or  within  the  corolla 
and  stamens ;  213.  In  a  capitulum  or 
head  of  flowers  it  is  the  central  part 
of  the  cluster,  or  the  whole  of  it  as 
opposed  to  a  border  or  ray.  It  is  the 
face  or  surface  of  any  organ,  such  as 
a  leaf-blade,  as  opposed  to  the  mar- 
gin. In  vegetable  anatomy,  certain 
round  spots  or  markings  on  cell-walls 
are  termed  discs. 

Discolor.  When  the  two  faces  of  a  leaf, 
&c.,  are  unlike  in  color. 

Discrete  (-etus).  Separate ;  not  coales- 
ced. 


408 


GLOSSAKY. 


Disepalous  (^us).    Two-sepaled;  244. 

Disk-flowert.  Those  belonging  to  the 
disk,  or  body,  and  not  to  the  margin 
or  ray  of  a  capitulum. 

Dissected  (-us).  Deeply  cut  or  divided 
into  numerous  segments. 

Dissepiment  (-entum).  A  partition  in  an 
ovary  or  pericarp ;  264. 

Dissilient  (-ens).  Bursting  asunder  or 
in  pieces. 

Distichous  (-us).    Disposed  in  two  ver- 

,    tical  ranks ;   122. 

Distinct  (Distinctus).  Separate  from; 
not  united. 

Distractile  (-His).    Carried  widely  apart. 

Dithecous  (-us).  Of  two  thecae,  or  cells, 
as  are  most  anthers ;  254. 

Diurnal.  Daily  ;  occurring  in  the  day; 
sometimes  used  for  ephemeral. 

Divaricate  (-atus).  Extremely  divergent. 

Divergent,  Diverging  (-ens).  Inclining 
away  from  each  other. 

Divided  (Divisus).  Where  lobing  or 
segmentation  extends  to  the  base ;  98. 

Dodeca.    Greek  for  twelve.    Used  in 

Dodecagynia.  Linnaean  order  with  flow- 
ers. 

Dodecdgynous.  Having  twelve  styles  or 
distinct  carpels. 

Dodecdmerous  (-its).  Of  twelve  parts  in 
the  circle. 

Dodecandria.  A  Linnsean  class  (334) 
with  the  flowers. 

Dodecdndrous.  Having  twelve  (or  from 
12  to  19)  stamens;  249. 

Dodrantdlis.  A  span  (about  nine  inches) 
long. 

Dolabriform  (-ormis).  Axe-shaped  or 
hatchet-shaped. 

Dorsal  (-alis).  Relating  to  the  dorsum 
or  back. 

Dorsal  Suture.  That  which  answers  to 
the  midrib  of  a  carpel ;  261. 

Dorsiferous.    Borne  on  the  back. 

Double.  Has  a  technical  use  when  a 
flower  is  said  to  be  "double;"  this 
denoting  one  in  which  the  leaves  of 
the  flowe?  are  monstrously  increased 
mostly  at  the  expense  of  the  essential 
organs. 

Downy.  Pubescent  with  fine  and  soft 
hairs.  Loosely  synonymous  with  soft- 
pubescent,  tomentose,  &c. 

Drepdnium.  A  sickle-shaped  cyme ;  156. 

Drupaceous  (-eus).  Resembling  or  relat- 
ing to  a  drupe. 

Drupe  (Drupa).    A  stone  fruit;  297. 

Drupelet,  Drupel  (Drupeola).  A  dimin- 
utive drupe ;  297. 


Drupetum.  An  aggregation  of  drupes; 
300. 

Duct.  In  vegetable  anatomy,  an  elon- 
gated cell  or  tubular  vessel,  found  espe- 
cially in  the  woody  (nbro-vascular) 
parts  of  plants. 

Dumetose  (-osus).  Pertaining  to  Dume- 
tum,  a  thicket,  or  Dumus,  a  bush. 

Dumose  (-osus).  Bushy,  or  relating  to 
bushes. 

Duplo.  Twice  as  many.  In  Greek  com- 
pounds, Diplo. 

Duramen.  The  heart-wood  of  an  exo- 
genous stem  ;  80. 

Dwarf.  Of  small  size  or  height  com- 
pared with  its  relatives. 

Dyclesium.    See  Diclesium. 


E-  or  Ex-  As  a  prefix  to  Latin  words, 
carries  a  privative  meaning,  as  Ecos- 
tate,  without  ribs,  Exalbuminous,  with- 
out albumen. 

Eared.     Same  as  Auriculate. 

Ebracteate,  Ebracteolate  (-atus).  Desti- 
tute of  bracts  or  bractlets. 

Eburneus.     Ivory-white. 

Ecdlcarate  (-atus).     Spurless. 

Echinate  (-atus).  Beset  with  prickles, 
like  a  hedgehog. 

Echinulate  (-atus).  Beset  with  diminu- 
tive prickles. 

Edentate  (-atus).     Toothless. 

Effete  (-etus),  or  Effcetus.  Past  bearing; 
functionless  from  age. 

Efflorescence  (-entia).  The  time  or  state 
of  blossoming ;  anthesis. 

Effuse  (-usus).  Very  loosely  spreading, 
more  so  than  diffuse. 

Eglandulose  (-osus).   Destitute  of  glands. 

Egg-shaped.    See  Ovate. 

Eldter.  One  of  the  spiral  or  spirally-- 
marked threads  in  the  spore-cases  of 
certain  Hepatic*. 

Eldtus.    Tall  or  lofty. 

Eleutheros.  In  Greek  compounds,  sep- 
arate or  distinct. 

Eleutheropetalous  (-us).  Same  as  Chori- 
petalous  or  Polypetalous ;  245. 

Ellipsoidal  (-eus).  An  elliptical  solid; 
sometimes  used  for  nearly  elliptical. 

Elliptical  (-us).  In  the  form  of  an  el- 
lipse. Oval  or  oblong  with  regularly 
rounded  ends;  95. 

Emarcidus.  Flaccid  or  withered. 

Emarginate  (-atus).  With  a  notch  cut 
out  of  the  margin ;  or,  as  usually  ap- 
plied, out  of  the  extremity ;  97. 

Embracing.    Clasping  by  the  base. 


GLOSSARY. 


409 


Embryo  or  Embryon.    The  rudimentary 

plantlet  formed  in  a  seed  ;  9,  311. 
Embryonal.     Relating  to   the  embryo; 

as  Embryonal  Vesicle ;  284. 
Embryo-sac.     The  cell  in  the  ovule  in 

which  the  embryo  is  formed ;  283. 
Embryogeny.     Embryo-formation. 
Emersed  (Emersus).    Raised  above  and 

out  of  the  water. 
Enantiobldstus.    With  embryo  at  the  end 

of  the  seed  diametrically  opposite  the 

hilum. 
Enation.    Having  outgrowths  from  the 

surface,  &c.;  179. 
Endeca.    In  Greek  compounds,  eleven; 

as  in 
Endecdndrous,    Endecdgynous.       With 

eleven  stamens  or  eleven  styles,  &c. 
Endemic.     Confined  geographically  to 

the  particular  region. 
£ndocarp  (-drpium).   The  inner  layer  of 
/  a  pericarp ;  288. 
Endockrome  (-oma).     Peculiar  coloring 

matter  in  cells;  especially  the  color- 
r  ing  matter  of  Algae. 
Endogens,     Endogenee.         Endogenous 

Plants;  70. 

Endogenous  structure,  70. 
Endopleura.     Inner  seed-coat ;  306. 
Endophlceum.     Inner  bark ;  77. 
Endorhizal  (-us).      Said  of  an  embryo 

which    has   the   radicle   sheathed   by 

the   cotyledon   or    plumule    wrapped 

around  it  in   many  Monocotyledons ; 

hence 

Endorhizce.     Synonym   of  Monocotyle- 
,  dones. 
Endosperm  (-ermium).    Synonym  of  the 

albumen  of  a  seed ;  or  the  inner  albu- 
,  men  ;  14,  310. 
Endostome  (-oma).     The  foramen  of  the 

inner  coat  of  an  ovule ;  277. 
Endothecium.    Inner  lining  of  the  cell  of 

an  anther. 
Enertis,  Enervius.    Nerveless;   no  ribs 

or  veins  visible. 
Ennea.     In    Greek  compounds,  nine ; 

as  in 
Enneagynia.    A  Linnaean  ordinal  name, 

and 
Ennedgynous.   With  nine  separate  styles 

or  carpels ;  337. 
Enneandria.    Linnaean  class,  and  Enne- 

androus,  with  nine  stamens ;  249,  334. 
Enodal  (Enodis).    Without  a  node. 
Ensdtus.     Same  as  Ensiform. 
Ensiform  (-ormis).    Sword-shaped ;  *.  e. 

like  a  broad  sword,  or  the  leaf  of  an 

Iris. 


Entire.  Without  toothing  or  dmsion,- 
the  margin  whole  and  even ;  97. 

Entomophilous.  Said  of  flowers  which 
are  habitually  fecundated  by  pollen 
,  carried  by  insects  ;  217,  218. 

Entophytes  (Entophyta).  Plants  grow- 
ing in  or  out  of  other  plants,  as  cer- 
tain Fungi,  &c. ;  4. 

Ephemeral.    Lasting  only  for  one  day. 

Epi.    In  Greek  compounds,  upon. 

Epiblast  (-us).  Name  sometimes  givea 
to  the  first  (and  an  undeveloping)  leaf 
of  the  plumule  of  the  embryo  of  grasses 
and  grain. 

Epiblastema.  A  superficial  outgrowth 
from  leaves,  &c. ;  210. 

Epicalyx.  Name  sometimes  given  to 
an  involucel  resembling  an  accessory 
calyx. 

Epicarp  (Epicarpium).  The  external 
layer  of  a  pericarp ;  288. 

Epichilium.  The  terminal  portion  of 
the  labellum  of  an  Orchid,  when  this 
is  of  two  parts. 

Epiclinal  (-us).     Upon  a  torus. 

EpicdroUine.     Upon  a  corolla. 

Epidermis.   The  skin  of  a  plant;  76,  89. 

Epigceous  (-eus).  Growing  on  or  out  of 
the  ground. 

Epigynous  (-us).  Literally  on  the  pistil; 
meaning  on  the  ovary,  or  seemingly 
so;  183. 

Epipetalous  (-us).  Borne  on  (adnate  to) 
the  petals ;  also  used  in  the  sense  of 
placed  before  the  petals. 

Epiphloeum.  The  outermost  or  corky 
bark;  76. 

Epiphyllous  (-us).    Growing  on  leaves. 

Epiphytal.    Pertaining  to 

Epiphytes  (Epiphyta).  Plants  growing 
on  other  plants  by  way  of  attachment, 
but  not  parasitic ;  air-plants ;  35. 

Epipteraus  (-us).  Winged  or  wing- 
bearing  at  summit. 

Episepalous.  On  the  sepals;  also  used 
in  the  sense  of  standing  before  a 
,  sepal. 

Episperm  (Epispermium).  The  coat  or 
f  outer  coat  of  a  seed ;  305. 

Epitropovs  (-us).  Name  (by  Agardh)  of 
an  anatropous  ovule  with  rhaphe 
averse  when  ascending,  adverse  when 
suspended;  282. 

Equal  (jEqualis).  Alike  as  to  length  or 
,  number,  &c.,  as  the  case  may  be. 

Equitant  (-ans).  Riding;  folded  around, 
as  if  straddling  over;  108,  138. 

Erect  (Erectus).  Standing  upright, 
mostly  in  relation  to  the  ground, 


410 


GLOSSARY. 


sometimes  when  perpendicular  to  the 
surface  of  attachment. 

Erion.  Greek  for  wool;  used  in  com- 
pound words,  as 

Erianthus.    With  woolly  flowers. 

Eridphorus.    Wool-bearing. 

Eriophyllus.     Woolly-leaved. 

Erostrate  (Erostris).    Beakless. 

Erose  (Erosus).  As  if  gnawed ;  applied 
to  an  irregularly  toothed  or  eroded 
margin. 

Erythros.  Greek  for  red,  used  in  com- 
pound terms  from  the  Greek. 

Erythrdstomum.  Name  given  by  Des- 
vaux  for  such  an  aggregate  fruit  as 
a  raspberry;  300. 

Estivation.    See  ^Estivation. 

Etcerio.  Name  of  aggregate  fruits, 
especially  of  fleshy  ones,  such  as  a 
blackberry. 

Etiolated.    Blanched  by  darkness. 

Eu.  Prefixed  to  words  of  Greek  deri- 
vation denotes  very,  true,  or  much  so. 
Frequently  used  in  names  of  sections 
or  other  groups ;  357. 

Euphylla.   Foliage-leaves,  or  true  leaves. 

Eutropic  (-icus).  Name  suggested  for 
twining  "with  the  sun;"  51. 

Evalvular  (Evalvis).  Not  opening  by 
valves. 

Evergreen.  Bearing  green  foliage  all 
the  year  round. 

Evittatut.    Not  vitiate. 

Ex.  A  prefix  in  place  of  E  privative 
when  the  following  part  of  the  com- 
pound begins  with  a  vowel ;  as 

Exalbuminous  (-osus).  Destitute  of  albu- 
men ;  14,  309. 

Exdlate  (-atus).    Destitute  of  wing. 

Exanthemata.  Eruptive  excrescenses  on 
the  surface  of  leaves,  &c. ;  blotches. 

Exaristate  (-atus).  Destitute  of  an 
arista  or  awn. 

Exasperate  (-atus).  Rough  with  hard 
projecting  points. 

Excentric  (-icus).  Out  of  the  centre; 
one-sided. 

Excurrent  (-ens).  Running  through  to 
the  very  summit  or  beyond ;  48. 

Exiguus.     Small  or  mean. 

Exilis.    Lank  or  meagre. 

Kximius.  Distinguished,  as  for  size  or 
beauty. 

Exo-.  In  Greek  compounds,  external  or 
,  outward ;  as 

Exocarp  (Exocarpium).  The  outer  layer 
of  a  pericarp  ;  288. 

Exogenous.  Outside  growing,  as  the 
wood  of  Dicotyledons ;  69,  73. 


Exogens,  Exogenat.  Exogenous  Plants; 
69,  340. 

Exorhizce.  Name  equivalent  to  Exogen, 
from 

Exorhizal  (-us).  The  radicle  not  sheathed, 
so  the  primary  root  in  germination 
/  has  no  covering  to  break  through. 

Exostome  (Exostoma).  The  foramen  of 
the  outer  coat  of  the  ovule ;  277. 

Exothecium.  The  outer  wall  of  an 
f  anther. 

Explanate  (  atus).    Spread  out  flat. 

Exsert,  Exserted  (Exsertus).  Protruding 
beyond  or  out  of,  as  stamens  beyond 
,  the  corolla. 

Exstipulate  (-atus).  Destitute  of  stipules. 

Exterior.  External  in  the  sense  of  outer. 
But  also  in  the  flower  sometimes  used 
in  the  sense  of  anterior. 

Extine.    Outer  coat  of  a  pollen-grain. 

Extra-axillary.  Beyond  or  out  of  the  axil. 

Extrorse  (Extrorsus,  Extrorsum).  Di- 
rected outward ;  253. 

Eye.  A  gardener's  name  for  an  unde- 
veloped bud. 

Fades.    Face ;  the  general  aspect. 

Falcate  (-atus),  and  Falciform  (-ormis). 
Scythe-shaped  or  sickle-shaped ;  plain 
and  curved,  with  the  edges  parallel. 

Family.  In  botany,  synonymous  with 
Order;  325. 

Fan-shaped.    See  Flabelliform. 

Farina.    Starch. 

Farinaceous  (-eus).  Of  the  nature  of 
starch,  or  containing  it. 

Farinose  (-osus).  Covered  with  a  meal- 
like  powder. 

Fasciate  (-atus).  Said  of  monstrous  ex- 
pansions of  stems,  giving  the  appear- 
ance as  of  several  stems  coalescent  in 
one  plane. 

Fascicle  (-icula).  A  close  cluster  or 
bundle,  whether  of  flowers,  stalks, 
roots,  or  leaves;  147,  153. 

Fascicled  ( Fasciculatus,  Fascicularit). 
In  a  fascicle;  131. 

Fastigiate  (-atus).  Said  of  branches 
when  parallel,  clustered,  and  erect 

Faux,  pi.  fauces.  The  gorge  or  throat 
of  a  gamophyllous  calyx  or  corolla; 
either  at  the  orifice,  or  a  portion 
between  the  limb  and  the  proper 
tube;  246. 

Fdveolate  (-atus),  Favose  (-osus).  Honey- 
combed ;  same  as  Alveolate. 

Feather-veined.  Having  veins  all  pro- 
ceeding from  the  sides  of  a  midrib. 

Feathery.    See  Plumose. 


GLOSSARY. 


411 


Fecula  or  Fascula.  Starch-like  matter. 
Applied  to  a  pistillate  flower,  or  to  a 
plant  producing  only  such  flowers. 

Fenestrate  (-atus),  Fenestralis.  Pierced 
with  large  holes,  like  windows. 

Ferrugineous  or  Ferruginous  (Ferrv- 
gineus).  Colored  to  imitate  iron-rust. 

Fertile  (-His).  Fruitful,  fruiting,  or  ca- 
pable of  producing  fruit ;  as  a  fertile 
flower  is  one  provided  with  a  well- 
formed  pistil ;  191.  In  English  descrip- 
tions, Flower  fertile  usually  means  a 
pistillate  or  female  flower.  Stamens 
or  anthers  are  also  said  to  be  fertile 
when  polliniferous  and  capable  of  fer- 
tilizing. 

Fertilization.  Synonym  of  fecundation, 
as  of  the  ovule  by  pollen;  215. 

Fibre  (Fibra).  Any  fine  filament;  the 
elementary  components  of  wood,  &c. ; 
delicate  roots,  &c. 

Fibril  (-ilia).    A  diminutive  fibre. 

Fibrillate  (-atus),  Fibrittose  (-osus). 
Furnished  or  abounding  with  fibres 
or  fibrils. 

Fibrous,  Fibrose  (-osus).  Composed  or 
of  the  nature  of  fibres. 

Fibro-vascular.  Consisting  of  woody 
fibres  and  ducts. 

Fiddle-shaped.  Obovate  and  with  a 
sinus  or  contraction  on  each  side. 

Fidus.  A  Latin  termination  for  cleft  or 
lobed. 

Filament  (-entum).  The  stalk  or  sup- 
port of  an  anther;  ,165,  251.  Also  any 
fibre-shaped  or  thread-like  body. 

Filamentous,  Filamentose  (-osus).  Com- 
posed of  threads  or  filaments. 

Filicology.  The  botany  of  Ferns :  re- 
placed by  Pteridology. 

Filiform (-ormis)  .  Thread-shaped;  long, 
slender,  and  terete. 

Filipendulous  (-us).  Hanging  from  a 
thread. 

Fimbria.    A  fringe,  or  dissected  border. 

Fimbriate  (-atus).  Fringed ;  bordered 
by  slender  processes  or  marginal  ap- 
pendages. 

Fimbrillate  (-atus),  Fimbrilliferous  (-as). 
Bearing  Fimbrillce  or  diminutive  fringe. 

Fingered.    See  Digitate. 

Fissiparous.  Multiplying  by  the  divi- 
sion of  one  body  into  two,  and  so  on. 

Fissus.    Split  or  cleft.    See  Fidus. 

Fistular,  Fistulose  (-osus).  Hollow 
through  the  whole  length,  as  the  leaf 
and  stem  of  an  Onion. 

Fldbellate  (-atus),  Flabelliform  (-ormis). 
Fan-shaped;  much  dilated  from  a 


wedge-shaped  base,  and  the  broader 
end  rounded. 

Flabellinerved.  With  radiating  straight 
nerves;  92. 

Flagellate  (-atus),  Flagellaris.  Produc- 
ing filiform  runners  (Flagella),  or 
runner-like  branches. 

Flagelliform  (-ormis).  Runner-like ; 
long,  slender,  and  supple  like  a  whip- 
lash or  Flagellum ;  53. 

Flammeus.    Flame-colored. 

Flavescent  (-ens).  Yellowish  or  pale 
yellow. 

Flavus.    Pale  yellow  or  ochre-yellow. 

Fleshy.  Succulent;  of  the  consistence 
of  flesh. 

Flexuous,  Flexuose  (-osus).  Zigzag; 
bent  alternately  in  opposite  direc- 
tions. 

Floating.  Borne  on  the  surface  of  water. 

Floccose  (-osus).  Bearing  or  clothed  with 
locks  of  soft  hairs  or  wool  (Jlocci). 

Flocculent.    Diminutive  of  floccose. 

Flora  (Goddess  of  flowers).  The  aggre- 
gate of  the  plants  of  a  country  or  dis- 
trict; or  the  name  of  a  work  which 
systematically  describes  them ;  369. 

Floral.    Belonging  to  the  flower. 

Floral  Envelopes.     Flower-leaves ;  164. 

Floret.   A  small  flower,  one  of  a  cluster. 

Floribundus.    Abundantly  floriferous. 

Floriferous(Florifer,Floriferus).  Bear- 
ing flowers. 

Florula.  A  small  Flora;  the  Flora  of 
a  restricted  district. 

Flos.  Latin  for  flower.  —  Flosplenus.  A 
"  double  "  flower ;  that  is,  one  in  which 
petals  are  increased  abnormally,  com- 
monly at  the  expense  of  the  androe- 
cium  or  the  gyncecium  also ;  171. 

Flosculus.    Latin  for  floret. 

Flower.  The  whole  reproductive  appa- 
ratus in  a  phaenogamous  plant ;  163. 

Flower-bud.  An  unexpanded  blossom 
or  undeveloped  cluster;  40. 

Flowering  Plants,  3,  344. 

Flowerless  Plants,  3,  344. 

Fluitans.    Floating. 

Fluviatile,  Fluvidtilis.  Belonging  to  a 
river  or  running  water. 

Fly-traps,  113. 

Fcemineus.  Feminine  or  female  flower, 
plant,  &c. ;  191. 

Foliaceous  (-eus).  Leaf-like  in  texture 
or  appearance ;  or  bearing  leaves. 

Foliar  (Foliaris).     Relating  to  leavea. 

Foliation  (Foliatio).     Leafing  out. 

Foliate  (-atus).  Having  leaves.  With 
Latin  numerical  prefix,  bifoliate,  tr»- 


412 


GLOSSARY. 


foliate,  and  so  on,  according  to  the 
number. 

Foliiform  (-ormis).     Leaf-shaped. 

Foliolate  (-atus).  Having  leaflets :  their 
number  may  be  indicated  by  Latin 
numerals,  as  bifoliolate,  trifoliolate, 
&c.;  102. 

Foliolum.    A  leaflet;  102. 

Foliose.     Bearing  numerous  leaves. 

Folium.    Latin  for  leaf ;  85. 

Follicetum.  A  whorl  or  aggregation  of 
follicles;  300. 

Follicle  (-iculus).  Fruit  of  a  single  carpel 
dehiscent  by  one  (the  ventral)  suture; 
292.  Name  of  the  earlier  botanists 
ibr  any  kind  of  capsular  fruit. 

Follicular  (-art's).  Pertaining  to  or  like 
a  follicle. 

Foot-stalk.  Petiole,  85 ;  or  Peduncle,  143. 

Foramen.  An  aperture  of  any  kind; 
specially  that  of  the  coat  of  the  ovule ; 
277. 

Foraminulose  (-osus).  Pierced  with  many 
small  holes. 

Forcipate  (-atus).  Like  forceps  or  pin- 
cers. 

Forked.   Divided  equally  into  branches. 

Fornicate  (-atus).  Arched  over,  as  by 
scales  (Fornices)  covering  the  throat 
of  the  corolla  of  Hound' s-tongue,  &c. 

Foveate  (-atus),  and  diminutive  Foveo- 
late.  Pitted;  impressed  with  shallow 
depressions  or  pits,  Fovece. 

Fovilke.  Minute  granules  in  a  liquid, 
in  the  protoplasm  of  the  pollen-grain, 
&c.;  258. 

Free.  Not  adnate  to  other  organs. 
Sometimes  used  in  the  sense  of  dis- 
tinct, i.  e.  unconnected  with  others  of 
the  same  sort. 

Fringed.    See  Fimbriate. 

Frond  (Frons).  An  old  name  for  leaf: 
employed  mainly  for  the  leaf  of  Ferns 
and  other  Cryptogamia,  and  certain 
Phaenogamous  plants  which  serve  for 
fructification  as  well  as  foliage;  also 
for  the  peculiar  foliage  of  Palms;  67. 

Frondescence  (-entia).  The  act  of  leaf- 
ing. Has  also  been  emplor^d  to 
express  the  metamorphosis  c*  floral 
organs  into  foliage-leaves ;  174. 

Frondose  (-osus).  Sometimes  used  in 
the  sense  of  leafy ;  also  frond-like,  or 
bearing  fronds. 

Fructification.  The  act  or  the  organs 
of  fruiting  or  reproduction  through 
flower  and  seed,  or  their  analogues. 

Fructus.    Latin  for  Fruit. 

Fruit.    The  immediate  product  of  fruc- 


tification ;  in  phsenogamous  plants, 
the  seed-vessel  and  contents,  along 
with  all  intimately  connected  acces- 
sory parts;  286. 

Fruit-dots  in  Ferns.     See  Sorus. 

Frumentaceous.  Relating  to  grain  (Fru- 
mentum). 

Frustulose  (-osus).  Consisting  of  similar 
pieces  or  Frustules  (frustula). 

Frutex.    A  shrub. 

Frutescent  (-ens).  Shrubby,  or  becom- 
ing so. 

Fruticose  (-osus).  Shrubby,  or  relating 
to  shrubs. 

Fruticulose  (-osus).  Relating  to  a  di- 
minutive shrubby  plant. 

Fruticulus.  A  minute  or  low  shrubby 
plant. 

Fugacious.  Falling  or  fading  very  early ; 
lasting  a  very  short  time. 

Fulvous  (-us).  Tawny;  orange-yellow 
and  gray  mixed. 

Fulcra.  Accessory  organs,  such  as  ten- 
drils, stipules,  spines,  and  the  like. 

Fulcrate  (-atus).  Propped,  supported 
by,  or  provided  with  accessory  organs. 

Fuliginous  (-osus).     Sooty-brown. 

Fungiform  (-ormis  and  Fungittiformis). 
Mushroom-shaped. 

Funaose  (-osus).  Spongy  in  texture; 
fungus-like. 

Funicule,  Funiculus.  The  stalk  of  an 
ovule  or  seed;  276. 

Funnelform,  Funnel-shaped;  249.  See 
Infundibuliform. 

Furcate  (-atus).  Forked;  or  divergently 
branched. 

Furcellatus.    Diminutively  forked. 

Furfuraceous  (-eus).  Scurfy;  covered 
with  bran-like  scales  or  powder. 

Furrowed.    See  Sulcate. 

Fuscous  (-us).    Grayish-brown  in  hue. 

Fusiform  (-ormis).  Spindle-shaped ; 
terete  and  tapering  gradually  to  each 
end;  31. 


Gdlbulus.  The  peculiar  strobile  of  Cy- 
press and  Juniper,  composed  of  up- 
wardly thickened  or  fleshy  scales; 
303. 

Galea.  A  helmet;  name  given,  from 
its  shape,  to  the  upper  sepal  of  Aconi- 
tum,  and  the  upper  lip  of  certain 
forms  of  bilabiate  corolla;  247. 

Galeate  (-atus).  Having  a  galea;  hel- 
met-shaped; 247. 

Gamo-.  In  Greek  compounds,  denotes 
union  by  the  edges  or  coalescence. 


GLOSSARY. 


413 


Gamopetalous.    A  corolla  of  coalescent 
petals ;  formerly  Monopetalous  ;  244. 
Gamophyllous  (-us).      Composed  of  coa- 
lescent leaves. 
Gamosepalous.     A  calyx   of   coalescent 

sepals;  244. 

Geitonogamy.     Fecundation  of   a  pistil 
by  pollen   of   another   flower  of  the 
same  plant;  216. 
Geminate  (-atus).     Twin ;  in  pairs ;  two 

side  by  side. 

Gemma.     A  bud,  specially  a  leaf-bud. 
Gemination.     Budding-growth;    or  the 

disposition  of  buds. 

Gemmule   ( Gemmula).      Diminutive   of 
gemma ;  minute  and  simple  buds  or 
bodies   analogous   to  buds;   also   sy- 
nonym of  Plumule.  For  a  time  used  by 
Endlicher  and  others  for  the  ovule. 
Genera.    Plural  of  Genus ;  323. 
General  (-alis).     Opposed  to  partial ;  as 

general  involucre. 
Generic.     Relating  to  genus. 
Genetic.  Genealogical ;  that  which  comes 

by  inheritance. 
Geniculate  (-atus).     Bent  abruptly,  like 

a  knee. 
Genitalia.    The  stamens  and  pistils  or 

their  analogues. 

Genus.     Kind  or  group  superior  to  spe- 
cies, and  which  with  the  species  gives 
the  name  to  the  plant ;  323. 
Geoblast  (-astus).     A  plumule  which  in 
germination  rises  from  underground, 
such  as  that  of  the  Pea. 
Germ.  A  growing  point  or  initial  growth, 
as  of  a  bud;  or  the  Embryo;  311.    Or 
in  the  sense  of 
Germen.     The    Linnsean   name  of  the 

ovary;  166. 

Germination  (-atio).     The  act  of  devel- 
opment of  the  embryo  of  a  seed  into  a 
plant. 
Gerontogceous  (-ceus).     Belonging  to  the 

Old  World. 

Gibbous,  Gibbose  (-osus).  Swelling  out 
on  one  side  into  a  gibber  or  gibber- 
osity. 

Giganteus.    Of  unusual  height. 
Gilvus.   Dirty  yellow  with  a  tinge  of  red. 
Glabrous  ( Glaber).    Smooth  in  the  sense 

of  not  pubescent  or  hairy. 
Glabrate  (-atus).     Somewhat  glabrous, 

or  becoming  glabrous. 
Glabriusculus.     Almost    but   not  quite 

glabrous. 

Gladiate  (-atus).  Sword-shaped  ;  in  the 
form  of  a  sword-blade,  whether  straight 
or  somewhat  curved-  See  Ensiform. 


Gland  for  Glans.    An    acorn  and  the 

like;  296. 

Gland  ( Glandula).  A  definite  secreting 
surface  or  structure  on  the  surface  of 
any  part  of  a  plant,  or  partly  imbedded 
in  it,  extended  to  any  protuberance 
or  structure  of  similar  nature  which 
may  not  secrete. 

Glandular,  Glandulose  (-osus).     Bearing 
glands  or  having  the  nature  of  glands. 
Glanduliferous  (-us).     Gland-bearing. 
Glareosus.    Growing  in  gravel. 
Glaucescent  (-ens).    Verging  upon  or  be- 
coming glaucous. 

Glaucous   (-us).     Covered   or  whitened 
with  a  bloom,  like  that  on  a  Cabbage- 
leaf. 
Globose  (-osus).    Having  or  approaching 

a  spherical  form. 

Globular  (-aris),Globulose  (-osus).  Some- 
what or  nearly  globose. 
Glochideous,  Glochidiate  (-atus).    When 
bristles  and  the  like  are  barbed  at  tip. 
Glockis.    A  barb. 
Glomerate  (-atus).   Compactly  clustered, 

especially  into  a 

Glomerule '  ( Glomerulus,    Glomus).       A 
cyme  condensed  into  a  head  or  capi- 
tate cluster ;  152. 
Glossology,  3,  359. 

Glumaceous  (-eus).  Pertaining  to  or  re- 
sembling glumes. 

Glume,  Gluma.     One  of  the   chaff-like 
bracts  of  the  inflorescence  of  Grasses 
and  their  relatives  ;  143. 
Glumella.      Diminutive  of  gluma  ;   an 

inner  or  secondary  glume. 
Glutinous  ( -osus).    Covered  with  a  sticky 

exudation. 

G6nophore(Gonophorum).  A  stipe  which 
elevates  both  stamens  and  pistil;  212. 
Gossypine  (-inus).     Cottony;  flocculent. 
Grdcilis.    Slender. 
Grain.    See  Caryopsis. 
Gramineous  (-eus).    Relating  to  grass  or 

grain-bearing  plants. 
Granular     (-am),     Granulose     (-osus). 
Composed  of  small  grains  or  Granules. 
Granulate  (-atus),  Granuliferus.     Bear. 

ing  grains  or  grain-like  bodies. 
Graveolens.  Unpleasantlystrong-scented. 
Griseus.    Gray  or  bluish-gray. 
Grumous    (Grumosus).      Consisting    of 

clustered  grains. 

Guttate  (-atus).    Spotted  as  if  by  drops. 
Gymndnthous  (-us).    Naked  flowered. 
Gymnos.   Greek  for  naked ;  used  in  com- 
pounds such  as 
Gymnocdrpous  (-us).     Naked-fruited. 


414 


GLOSSARY. 


Gymnospermia.  A  Linnsean  artificial 
order  of  Didynamia,  in  which  the 
nutlets  resulting  from  four  divisions 
of  aa  ovary  were  taken  for  naked 
seeds;  337. 

Gymnosperms,  Gymnospermce.  A  sub- 
class of  naked-seeded  plants ;  268,  344. 

Gymnospermous  (-us).  Naked-seeded,  as 
opposed  to  Angiospermous. 

Gynandria.  A  Linnaean  class,  character- 
ized by  the  flower  being 

Gynandrous.  Stamens  borne  on  (adnate 
to)  the  pistil,  even  to  the  style  or 
stigma;  251,  335. 

Gynobase  ( Gynobasis).  An  enlargement 
or  production  of  the  torus  on  which 
the  gyncecium  rests  or  is  somewhat 
elevated;  212. 

Gyno-dicecious.  Dioecious  with  some 
flowers  hermaphrodite  and  others  pis- 
tillate only;  191. 

Gyncecium.  The  pistil  or  collective  pis- 
tils of  a  flower ;  the  female  portion  of 
a  flower  as  a  whole ;  165. 

Gynophore  (Gyndphorum).  The  stipe  of 
a  pistil ;  212. 

Gynostegium.  A  sheath  or  covering  of 
the  gynoecium,  of  whatever  nature. 

Gynostemium.  The  column  of  an  Orchid, 
consisting  of  andrcecium  and  summit 
of  the  gynoecium  combined. 

Gyrate  (-atus).  Curved  into  a  circle,  or 
taking  a  circular  course. 

Gyrose  (-osus).  Curved  backward  and 
forward  in  turns. 


Habit  (Habitus).  The  general  appear- 
ance of  a  plant. 

Habitat.  Habitation;  the  geographical 
limits  or  station;  366. 

Hcematitic  (-icus).    Brown-red. 

Hairt.  Outgrowths  of  the  epidermis, 
consisting  of  single  elongated  cells,  or 
of  a  row  of  cells. 

Hairy.  Descriptively  applied  to  pilosity 
or  pubescence,  in  which  the  hairs  are 
separately  distinguishable. 

Halbert-  or  Halberd-shaped.  See  Has- 
tate. 

Halved.  See  Dimidiate;  with  one  half 
absent  or  appearing  to  be  so. 

Hamate  (-atus).     Hooked  at  the  tip. 

Hdmulate  or  Hamulose  (-osus).  Dimin- 
utive of  Hamate. 

Haplos.  In  Greek  compounds,  simple 
or  simply,  as 

Haplopetalus  (-«#).  With  only  one  row 
of  petals. 


Haplostemonotu  (-us).  With  a  single 
series  of  stamens;  177. 

Hastate  (-atus),  Hastilis.  Halberd- 
shaped,  like  the  head  of  a  halberd, 
i.  e.  sagittate,  but  the  basal  lobes  di- 
rected outward  or  at  right  angles  to 
the  midrib  of  the  leaf;  96. 

Head.  The  form  of  inflorescence  termed 
Capitulum,  viz.  a  cluster  of  sessile 
flowers  on  a  very  short  axis  and  centri- 
petal in  evolution ;  147. 

Heart-shaped.  Ovate  with  a  sinus  at 
base;  96. 

Heart-wood.  The  older  and  matured 
wood  of  an  exogenous  stem ;  80. 

Hebetate  (-atus).  Having  a  dull  or  blunt 
and  soft  point. 

Helicoid  (-oideus),  Helicoidal.  Coiled  into 
a  helix,  or  like  a  snail-shell.  In  true 
helicoid  inflorescence,  the  flowers  are 
all  in  a  single  row;  155, 157. 

Helmet.     See  Galea. 

Helvolus.    Dull  and  grayish  yellow. 

Hemi.  Half  or  halved;  in  Greek  com- 
pounds, such  as 

Hemi-anatropous.     Half  anatropous. 

Hemicarp  (-arpium).  Half  or  one  carpel 
of  a  Cremocarp. 

Hemitropous  (-us).  Same  as  amphitro- 
pous  or  half  anatropous. 

Hepta.    The  Greek  numeral  seven. 

Heptagynia.  A  Linnaean  artificial  order, 
having  seven  styles  or  distinct  car- 
pels; 337. 

Heptdmerous  (-us).    Of  seven  members. 

Heptandria.  The  Linnsean  class  with 
seven  stamens;  334. 

Heptandrous.     Seven-stamened ;  249. 

Herb  (Herba).  A  plant  with  no  persist- 
ent woody  stem  above  ground ;  50. 

Herbaceous.  Of  the  texture,  color,  or 
other  characters  of  an  herb. 

Herbarium,  Herbal.  A  collection  of 
dried  specimens  of  plants,  systemati- 
cally arranged ;  380. 

Hercogamous  (-us).  Said  of  hermaphro- 
dite flowers  when  some  structural  ob- 
stacle prevents  autogamy. 

Hermaphrodite  (-itus).  Of  both  sexes; 
191. 

Hesperidium.  A  hard-rinded  berry,  like 
an  orange  and  lemon  ;  299. 

Heteracmy.  Synonym  of  Dichogamy  ; 
219. 

Heteros.  In  Greek  compounds,  denotes 
diverse  or  various,  as 

Heterocai-pous  (-us,  Heterocarpicus). 
Producing  more  than  one  kind  of 
fruit 


GLOSSARY. 


415 


Heterocephalous  (-««).  Bearing  two 
kinds  of  head  or  capitulum. 

Ifeterodite  (-itus).  Anomalous  in  forma- 
tion. 

Heterocline  (-inus).  Nearly  same  as 
Heterocephalous  ;  on  separate  recepta- 
cles. 

Heterodromous  (-us).  Spirals  of  changing 
direction. 

Heterdgamous  (-MS).  Bearing  two  kinds 
of  flowers. 

Heterogeneous.    Not  uniform  in  kind. 

Heterogone  or  Heterogoneus.  When  the 
flowers  are  dimorphous  or  trimor- 
phous  as  respects  relative  length,  &c., 
of  stamens  and  pistil;  225,  234. 

Heterostyled.    Same  as  Heterogone ;  234. 

Heterdmerous.  Of  members  not  corre- 
sponding in  number. 

Heterophyllous  (-us).  Having  leaves  of 
more  than  one  form. 

Heterotrdpous  (-us).  Turned  in  more 
than  one  direction,  or  in  an  unusual  di- 
rection ;  same  as  Amphitropous ;  279. 
(Also  used  by  Agardh  for  collateral 
ovules  turned  back  to  back ;  282. ) 

tfexa.  Greek  numeral  six ;  from  which 
is  formed 

Hexayynia.  Linnaean  artificial  order,  of 
flowers  with  six  styles  or  distinct  car- 
pels ;  337. 

Hexdgynous.  Having  the  character  of 
Hexagynia. 

Hexdmerous  (-us).  Of  six  members ;  176. 

Hexandria.  Linnaean  class  with  perfect 
flowers  of  six  stamens ;  334. 

Hexdndrous.    Having  six  stamens ;  249. 

Hexapetalous  (-us).    Having  six  petals. 

Hexaphyllous.    Six-leaved. 

Hexdpterous  (-us).     Six-winged. 

Hexasepalous.     Having  six  sepals. 

Hexaste~monous.     Having  six  stamens. 

Hibernaculum.    A  winter-bud;  40. 

Hidden.    Concealed  from  sight;  as 

Hidden-veined,  where  the  veins  are  in- 
visible, as  in  the  leaves  of  Pinks  and 
Houseleeks. 

Hiemal  (-alls).     Relating  to  winter 

Hilar  (Hilaris).  Belonging  to  the  hilum. 

Hllum.  The  scar  or  place  of  attachment 
of  the  seed ;  277,  305. 

Hippocrepiform  (-ormis).  Horseshoe- 
shaped. 

Hirsute  (-utus).  Pubescent  with  rather 
coarse  or  stiff  hairs. 

Hirtellous  (-us).     Minutely  hirsute. 

Hirtus.     Hairy,  nearly  same  as  Hirsute. 

Hispid  (-idus).  Beset  with  rigid  or 
bristly  hairs  or  with  bristles. 


Hispidulous  (-us).    Minutely  hispid. 

Hoary.  Grayish-white  with  a  fine  and 
close  pubescence.  See  Canescent. 

Holosericeous  (-eus).  Covered  with  fine 
and  silky  pubescence. 

Homocarpous  (-us).  With  fruit  all  of  one 
kind. 

Homodromous  (-us),  Homodromy.  With 
spirals  all  of  uniform  direction. 

Homogamous  (-us).  Bearing  one  kind 
of  flowers. 

Homogeneous.   All  of  one  nature  or  kind. 

Homdgonous  or  Homogme.  Homomor- 
phous  as  respects  the  stamens  and  pis- 
til ;  opposed  to  dimorphous ;  225. 

Homologue,  A  homologous  organ  or 
part. 

Homologous.  Of  one  name  or  type,  such 
as  leaves  and  parts  answering  morpho- 
logically to  leaves ;  6. 

Homomdilus.  Said  of  leaves  and  the  like 
which  are  all  turned  in  one  direction. 

Homomorphous  (-us).     All  of  one  form. 

Homostyled.     Same  as  Homogone. 

Homotropous  (-us).  Curved  or  turned  in 
one  direction ;  applied  also  to  the  em- 
bryo of  an  anatropous  seed,  with  rad- 
icle next  the  hilum ;  312. 

Hood.    See  Cucullus. 

Hooded.     Bearing  or  in  form  of  a  hood. 

Hornus,  Homotinus.  Of  the  present 
year. 

Horny.  Of  the  consistence  of  horn.  See 
Corneus. 

Hortensis,  Hortuldnus.  Pertaining  to 
the  garden. 

Hortussiccus.  Old  name  of  an  herbarium. 

Humi.     On  the  ground. 

Humifusus,  Humistrdtus.  Spread  over 
the  surface  of  the  ground. 

Humilif.     Low  of  stature. 

Hyaline  (-inus).  Transparent  or  trans- 
lucent. 

Hybrid.  A  mongrel,  or  cross-breed  of 
two  species ;  321. 

Hydrophytes  (Hydrophyta).  Water- 
plants. 

Hyemdlis.    See  Hiemalis. 

Hypanthium.  An  enlargement  or  other 
development  of  the  torus  under  the 
calyx;  214. 

Hypo.  In  Greek  compounds,  denotes 
under,  beneath,  lower. 

Hypanthodium.  Same  as  Syconium; 
149,  303. 

Hypochilium.  The  basal  portion  of  the 
labellum  of  an  Orchid. 

Hypocraterimorphous  (-us),  or  Hypocra- 
teriform,  but  the  latter  is  a  hybrid  of 


416 


GLOSSARY. 


Greek  and  Latin.  Salverform  or  sal- 
ver-shaped ;  that  is,  in  the  form  of  a 
salver  raised  on  a  central  support  or 
stem  beneath.  Said  of  a  corolla  and 
the  like  with  slender  tube  abruptly  ex- 
panded into  a  flat  limb;  248. 

Hypogceous  (-ceus).  Growing  or  remain- 
ing underground;  19. 

Hypogynous  (-us).  Under  or  free  from 
the  gynoecium  or  pistil ;  182. 

Hypophyllous  (-us).  Growing  on  the  un- 
der side  of  a  leaf. 

Hypophyllium.  An  abortive  leaf  or  scale 
under  another  leaf,  or  seeming  leaf,  as 
in  Asparagus  and  Ruscus. 

Hypsophylla.  Answers  to  the  German 
"  Hochblatter,"  or  high  leaves,  those 
of  the  inflorescence,  t.  e.  bracts  and 
the  like;  6. 

Hysteranthous  (-us).  With  leaves  pro- 
duced later  than  the  blossoms. 


Icosandria.  The  Linnaean  class  with 
twenty  stamens  (as  the  name  denotes) 
or  a  larger  number,  inserted  on  the 
calyx ;  334.  fcosandrous  is  the  corre- 
sponding adjective ;  249. 

Imberbis.     Not  bearded. 

imbricate  (-atus),  Imbricative.  Over- 
lapping so  as  to  "  break  joints,"  like 
tiles  or  shingles  on  a  roof ;  either  with 
parts  all  in  one  horizontal  row  or  cir- 
cle, as  in  the  aestivation  of  a  calyx  or 
corolla,  when  at  least  one  piece  must 
be  wholly  external  and  one  internal; 
or  with  the  tips  of  lower  parts  covering 
the  bases  of  higher  ones  in  a  succession 
of  rows  or  spiral  ranks ;  135. 

Immarginate  (-atus).  Not  margined  or 
bordered. 

Immersed  (-us).  Growing  wholly  under 
water. 

Impari-pinnate.  Pinnate  with  an  odd 
terminal  leaflet;  101. 

Incequilateral  (-alis).     Unequal-sided. 

Inanis.  Empty,  as  an  anther  containing 
no  pollen. 

Inappendiculate  (-atus).  Not  appen- 
daged. 

Incanescent.     Same  "as  Canescent. 

Incanus.     Hoary-white. 

Incarnate  (-atus).  Flesh-colored.  See 
Carneus. 

Incised  (-us).  Cut  irregularly  and 
sharply;  98. 

Included  (Inclusus).  When  the  part  in 
question  does  not  protrude  beyond  the 
surrounding  organ. 


Incomplete  (-«*).  Wanting  some  essen- 
tial component  part ;  190. 

Incrassate  (-atus).    Thickened. 

Incubous  (-us).  The  tip  of  one  leaf  or 
other  part  lying  flat  over  the  base  of 
the  next  above  it. 

Incumbent  (-ens).  Leaning  or  resting 
upon. 

Incumbent  Anther.  One  lying  against 
the  inner  face  of  filament ;  253. 

Incumbent  Cotyledons,  when  the  back 
of  one  lies  against  the  radicle;  313. 

Incurved  (-us).  Bending  from  without 
inward. 

Indejinite  (-itus).  Relates  usually  to 
number,  this  either  uncertain  or  too 
many  for  easy  counting. 

Indejinite  Growth,  49. 

Indejinite  Inflorescence,  same  as  Inde- 
terminate; 144. 

Indehiscent  (-ens).  Not  opening  by  valves, 
chinks,  or  along  regular  lines ;  288. 

Indeterminate.  Not  terminated  abso- 
lutely, as  the  inflorescence  in  which 
no  blossom  ends  the  axis  of  the  flower- 
cluster;  144,  146. 

Indigenous  (-^us).  Native  and  original 
to  the  country. 

Individuals,  315. 

Indivisus.  Undivided,  t.  e.  not  cleft, 
lobed,  or  parted. 

Indumentum.  Any  hairy  covering  or 
pubescence  which  forms  a  coating. 

Induplicate  (-atus).  With  edges  folded 
in  or  turned  inward. 

Indusium.  The  proper  (often  shield- 
shaped)  covering  of  the  sorus  or  fruit- 
cluster  of  a  Fern. 

Induviate  (-atus).  Clothed  with  with- 
ered parts  or  Induvice  (clothing). 

Inequilateral.     Unequal-sided;  106. 

Inermis.  Unarmed,  without  prickles, 
thorns,  &c. 

Inferior  (Inferus).  Said  of  one  organ 
when  below  another.  In  the  blossom 
also  in  the  sense  of  anterior ;  160.  An 
inferior  calyx  is  one  below  the  ovary, 
or  free ;  183.  An  inferior  ovary  is  one 
with  adnate  or  superior  calyx  ;  183. 

Inflated  (-atus).     Bladdery. 

Infiexed  (-us).  Bent  or  turned  abruptly 
inward. 

Inflorescence.  Mode  of  disposition  of 
flowers  ;  less  properly  used  for  a  flower- 
cluster  itself ;  141. 

Infra-axillary  (-aris).     Below  the  axil. 

Infundibuliform  (-ormis),  Infundibular 
(-aris).  Funnelform,  funnel-shaped; 
249. 


GLOSSARY. 


4T7 


Innate  (-atus).  Borne  on  the  apex  of 
the  supporting  part;  in  an  anther  the 
counterpart  of  adnate ;  252. 

Innovation  (-io).     A  new-formed  shoot. 

Inosculating.     Same  as  Anastomosing. 

Inseparate,  Inseparation.  Terms  pro- 
posed by  Masters  to  express  coales- 
cence; 181. 

Inserted  (-us).  Attached  to  or  growing 
out  of. 

Insertion  (-io).  Is  the  mode  or  place 
where  one  body  is  attached  to  that 
which  bears  it. 

Integer.  Entire  in  the  sense  of  un- 
divided, or  not  lobed  ;  97. 

Integerrimus.  Entire  in  the  sense  of 
quite  entire,  i.  e.  the  margin  without 
dentation;  97. 

Inter.  Between;  as  in  Intercellular, 
between  the  cells,  &c. 

Interfoliaceous  (-eus).  Between  the 
leaves  of  a  pair,  as  the  stipules  of 
many  Rubiacese. 

Internode  (-odium).  The  portion  of 
stem  between  two  nodes;  6. 

Interpetiolar  (-aris).  Between  the  pet- 
ioles. 

Interruptedly  pinnate.  Pinnate  with- 
out a  terminal  leaflet. 

Intine.  The  inner  coat  of  a  grain  of 
pollen. 

Intrafoliaceous  (-eus).  Within  or  be- 
fore a  leaf. 

Introfiexed  (-us).     Same  as  inflexed. 

Introrse  (-orsus).  Turned  inward  or 
toward  the  axis  ;  253. 

Introvenius.     Same  as  hidden-veined. 

Jntruse  (-u#us).  Pushed  or  projecting 
inward. 

Involucellate  (-atus).  Provided  with  a 
secondary  involucre  or 

Involucel  (-ellum).  An  inner  or  secon- 
dary involucre,  that  of  an  umbellet, 
&c.;  142. 

Involucrate  (-atus).  Provided  with  an 
involucre. 

Involucre  (Involucrum).  A  circle  of 
bracts  subtending  a  flower-cluster; 
,142. 

Involute  (-utus).     Rolled  inward;  133. 

Irregular  (-aris).  Exhibiting  a  want  of 
symmetry  in  form ;  184. 

Irregularity,  179,  184,  219. 

Inadelphous  (-us).  Equal  brotherhood,  as 
when  the  number  of  stamens  in  two 
phalanges  is  equal. 

l"6chrous.     All  of  one  color  or  hue. 

Isomerous  (-us).  The  members  of  suc- 
cessive circles  equal  in  number ;  175. 


Isostemonout  (^us).    The  stamens  just  as 

many  as  the  petals,  &c. ;  177. 
Isostemony,  196. 

Jointed.    See  Articulated. 

Juba.  A  loose  panicle,  with  axis  de- 
liquescent. 

Jugum,  pi.  juga.  A  pair  of  leaflets.  So 
pinnate  leaves  are  unijugate,  with  a 
single  pair  of  leaflets;  bijugate,  with 
two  pairs;  trijugate,  with  three  pairs 
or  juga,  &c.  Also  the  ridges  on  the 
fruit  of  Umbelliferae  are  termed  juga. 

Julus.    Same  as  Amentum  or  Catkin. 

Julaceous  (-eus).     Catkin-like,   Amen- 


Keel.  A  central  dorsal  ridge,  like  the 
keel  of  a  boat.  The  two  anterior 
petals  of  a  papilionaceous  corolla, 
which  are  united  into  a  body  shaped 
like  the  keel  or  the  prow  of  a  vessel ; 
185. 

Keeled.    Having  a  keel.     See  Carinate. 

Kernel.  The  nucleus  of  an  ovule,  or 
of  a  seed,  i.  e.  the  whole  body  within 
the  coats. 

Kermesinus.    Of  the  color  of  carmine. 

Key-fruit.    See  Samara;  294. 

Kidney-shaped.  Crescentic  with  the 
ends  rounded ;  very  oblately  cordate ; 
96. 

Kingdom,  325. 

Labellum.    One  of  the  petals  of  an  Or- 

chideous  flower,  which  is  unlike  the 

others. 
Labiate  (-us).    Lipped,  mostly  Bilabiate ; 

247. 
Labiatiflorous  (-us).      Said    of    certain 

Compositae  with  bilabiate  corollas. 
Labiose  (-osus).     Said  of  a  polypetalous 

corolla  which  has  the  appearance  of 

bilabiation. 
Labium.     See  Lip. 
Lacerate  (Lacerus).    Irregular  cleft  as 

if  torn  or  lacerated. 
Lacinia.    A  slash;  used  for  a  slender 

lobe. 
Laciniate    (-atus).      Slashed;    cut   into 

narrow  incisions. 

Lacinula.  A  diminutive  lacinia  or  nar- 
row lobe. 

Lactescent  (-ens).    Yielding  milky  juice. 
Lacteus.     Milk-white. 
Ldcunose  (-osus).      Abounding  in   pits, 

holes,  or  depressions  (lacuna). 
Lacustrine  (Lacustris).     Belonging  to  01 
i       living  in  lakes  or  ponds. 


418 


GLOSSARY. 


Lcevigate  (-atus).    Smooth  as  if  polished. 
Lcevis  (this  form,  and  not  levis,  has  al- 
ways been  used  in  botany).     Smooth 

in  the  sense  of  not  rough. 
Lageniform    (-ormis).      Shaped    like  a 

Florence  flask  or  a  gourd  (the  fruit  of 

Lagenaria). 

Lagdpus,     Hare-footed.     Densely  cov- 
ered with  long  hairs. 
Lamella.     A  thin  plate. 
Lidmellar  (-aris),  Lamellate  (-atus),  La- 

mellose   (-osus).     Composed    of  thin 

plates  or  lamellae. 
Lamina.    The  blade  or  expanded  part 

of  a  leaf,  &c.;  85,245. 
Lanate  (-atus),  Lanose  (-osus).    Bearing 

long    and    implexed    hairs    or    wool 

(lana). 
Lanceolate  (-atus).     Shaped  like  a  lance 

or  spear-head;  narrower  than  oblong, 

and  tapering  to  each  end,  or  at  least 

to  the  apex;  95. 
Lanuginous  (-osus).    Cottony  or  woolly ; 

clothed  with  soft  and  implexed  hairs 

or  down  (lanugo). 
Lnppaceus.     Like  a  bur  (lappa). 
Lasianthus.     Woolly-flowered. 
Latent.     Undeveloped  or  dormant,  as 

certain  buds ;  40. 
Lateral  (-alis).    Belonging  to  or  borne 

on  the  sides. 

Lateritious  (-ius).    Of  a  brick-red  color. 
Latex.    Proper  juice,  milky  juice,  and 

the  like. 
Laticiferous.    Containing  or  conveying 

latex. 

Latiseptus.     With  broad  partition. 
Lavender-color.     Pale  blue  with  some 

gray. 

Laxus.     Loose. 
Leaf.     The  principal  sort  of  appendage 

or  lateral  organ  borne  by  the  stem  or 

axis;  85. 

Leaf-blade.    The  lamina  of  a  leaf. 
Leaf-bud.     A  bud  which  develops  into 

a  leafy  branch  or  its  continuation ;  40. 
Leaflet.    A  blade  or  separate  division 

of  a  compound  leaf ;  100. 
Leaf-scar.     The  cicatrix  left  bv  the  ar- 
ticulation and  fall  of  a  leaf;  47. 
Leafstalk.     A  petiole  or  footstalk  to  a 

leaf-blade;  85,104. 
Leathery.     See  Coriaceous. 
Lecus.    A  synonym  for  Conn. 
Legume  (Legwnen).    The  seed-vessel  of 

Leguminosse,  a  carpel  which  normally 

dehisces  by  both  the  ventral  and  the 

dorsal  suture;  292. 

Perusing   to  a  legume, 


or  to  the  order  to  which  the  legume 
gives  its  name. 

Lenticels  (Lenticellce).  Lenticular  spots 
on  young  bark. 

Lenticular  (-aris).  Lens-shaped,  that  is 
the  shape  of  a  lentil  or  a  double-convex' 
lens. 

Lentiginosus.  Covered  with  minute  dots 
or  freckles. 

Lepal,  Lepalum.  A  made-up  word  to 
signify  a  stamen  transformed  into  a 
scale,  nectary,  &c. 

Lepicena.  Unused  name  for  a  glume  of 
Grasses. 

Lepidote(-otus).  Beset  with  small  scurfy 
scales. 

Lepis.    Greek  term  for  a  scale. 

Leptos.  Greek  for  slender ;  as  in  Lep- 
tophyllus,  slender-leaved. 

Leucos.    Greek  for  white ;  whence 

Leucanthus.       White-flowered. 

Leucophyllus.    White-leaved,  &c. 

Liber.  The  inner  and  often  fibrous 
bark ;  77,  81. 

Lid.     See  Operculum. 

Ligneous   (-eus),  Lignosus.     Woody. 

Ligule  (Ligula).  A  strap  or  strap- 
shaped  body,  such  the  principal  part 
of  a  ray- corolla  in  Compositse.  The 
thin  and  scarious  projection  from  the 
summit  of  the  sheath  of  the  leaf  of 
Grasses,  &c. ;  106.  Or  a  similar  out- 
growth of  the  inner  face  of  certain 
petals;  211. 

Ligulate  (-atus),  Liguliform  (-ormis). 
Furnished  with  a  ligule;  148,  247. 

Liguliflorous  (-us).  Said  of  the  head  of 
those  Compositse  which  contain  only 
ligulate  corollas. 

Liliaceous.    Lily-like;  246. 

Limb  ( Limbus).  A  border,  i.  e.  the  ex- 
panded part  of  a  gamophyllous  peri- 
anth, &c.,  as  distinguished  from  the 
tube  and  throat ;  245.  Sometimes  the 
term  is  applied  to  the  lamina  or  blade 
of  a  petal  or  a  leaf. 

Limbate  (-atus).     Bordered. 

Line  (Linea).  The  twelfth  part  of  an 
inch.  By  some  reduced  to  the  tenth 
of  an  inch ;  but  the  decimal  line  is  un- 
usual in  botanical  measurement. 

Linear  (-am).  Narrow,  several  times 
narrower  than  wide,  and  the  margins 
parallel;  95. 

Lineate  (-atus).     Marked  with  lines. 

Lineolate  (-atus).  Marked  with  fine  or 
obscure  lines. 

Lingweformis  or  Linguiformis,  also  Lin- 
ffvlate  (-aba). 


GLOSSAEY. 


419 


Lip.  One  of  the  two  divisions  of  a  bila- 
biate corolla  or  calyx,  t.  e.  of  a  gamo- 
phyllous  organ  which  is  cleft  into  an 
upper  (superior  or  posterior)  and  a 
lower  (inferior  or  anterior)  portion  or 
lip  (labium). 

Literal  or  Littoral  (-alis).  Belonging 
to  or  growing  on  the  seashore  or  river- 
shore. 

Livid  (-idus).    Pale  lead-colored. 

Lobe  (Lobus).  Any  division  of  an  organ ; 
or  specially  a  rounded  division  or  pro- 
jection; 98. 

Lobate  (-atus)  or  Lobed.  Divided  into 
or  bearing  lobes ;  98. 

Lobulate  (-atus).  Divided  into  small 
lobes.  Lobelets,  or  Lobules;  98. 

L6cellate  (-atus).  Divided  into  locelli; 
263. 

Locellus.  A  secondary  cell ;  as  where 
a  proper  cell  (loculus)  of  an  anther  or 
an  ovary  is  divided  by  a  partition  into 
two  cavities ;  251,  263. 

Loculament  (-entum).  Same  as  Loculus ; 
289. 

Locular  (-aria).  Celled;  as  bilocular, 
two-celled ;  trilocular,  three-celled, 
quadrilocular,  four-celled,  &c. 

Loculicidal  (-idus),  Loculicide.  Dehis- 
cent into  the  cell  or  cavity  of  a  peri- 
carp by  the  back,  t.  e.  through  a  dor- 
sal suture ;  289. 

Ldculus.  The  cell  or  cavity  in  an  orary 
or  an  anther. 

Loculote  (-osus).  Partitioned  off  into 
cells. 

L6custa.    Name  of  a  spikelet  in  Grasses. 

Lodicule,  Lodicula.  One  of  the  small 
scales  next  to  the  stamens  in  the  flower 
of  Grasses. 

Lomentaceous  (-eus).  Bearing  or  re- 
sembling a 

Loment  (Lomentum).  A  legume  which 
is  constricted  or  which  separates  into 
one-seeded  articulations ;  293. 

Lorate  (-atus).  Strap-shaped  or  thong- 
shaped;  same  as  much-elongated 
linear. 

Lucid  (-idus).     With  a  shining  surface. 

Lunate  (-atus).  Half  moon-shaped ; 
crescent-shaped. 

Lunulate  (-atus).    Diminutive  of  Lunate. 

Lupuline  (-inus).  Resembling  a  head  of 
Hops. 

Lurid  (-idus).     Dingy-brown. 

Lusus.  A  "sport"  or  variation  from 
seed  or  bud;  319. 

Luteolus.  Yellowish;  diminutive  of 
Luteua. 


Lutescent  (-ent).    Becoming  yellow,  or 

faintly  yellow. 
Luteus.    Latin  for  yellow. 
Lycotropous  (-us).    Said  of    an    ortho- 

tropous  ovule  when  bent  into  an  open 

curve  or  horseshoe  form. 
Lyrate  (-atus).     Lyre-shaped;  a  pinnat- 

ifid  form  with  terminal  lobe  large  and 

rounded  and  one  or  more  of  the  lower 

pairs  small;  hence  Lyrately pinnate ; 

101,  &c. 

Macros.  Greek  for  large  or  more  prop- 
erly long;  hence  Macranthus,  long- 
flowered  ;  Macrocephalus,  large- 
headed;  Macropodus,  long-footed,  or 
with  long  stalk,  &c. 

Mdcrospore.  The  larger  kind  of  spore 
in  Lycopodiaceae,  &c. 

Maculate  (-atus).  Spotted  or  blotched, 
i.  e.  with  macula. 

Malpighiaceous  Hairs.  Those  fixed  by 
the  middle  and  tapering  both  ways. 

Mdmillate  (-atus),  Mamillar  (-arts). 
Bearing  teat-shaped  processes. 

Mammceform  (-ormis).  Breast-shaped 
or  teat-shaped ;  conical  with  rounded 
apex. 

Mammosus.    Breast-shaped. 

Mancus.    Deficient  or  wanting. 

Mdnicate  (-atus).  Said  of  pubescence 
so  dense  and  interwoven  that  it  may 
be  stripped  off  like  a  sleeve. 

Marcescent  (-ens),  Marcidus.  Withering 
without  falling  off;  243. 

Marginate  (-atus).  Furnished  with  a 
margin  of  a  distinct  character  or  ap- 
pearance. 

Marginiddal.  Dehiscent  by  the  dis- 
junction of  the  united  margins  of  car- 
pels; 290. 

Marmoratus.  Marbled ;  traversed  by 
veins  or  shades  of  color. 

Maritime  (Maritimus).  Pertaining  to 
the  sea  or  seaeoast 

Mas,  Masculus,  Masculinus.  Belonging 
to  the  stamens,  or  staminate  plant,  or 
flower;  191,  &c. 

Masked.     See  Personate. 

Mealy.     See  Farinaceous. 

Medial,  Median  (Medidnus).  Belong- 
ing to  the  middle;  in  the  plane  of 
bract  and  axis ;  160. 

Medifixus.    Fixed  by  the  middle;  253. 

Medulla.     Pith;  75. 

Medullary.  Relating  to  the  pith.  Med- 
ullary Rays,  74 ;  Medullary  sheath,  75. 

Meiostemonous.  With  fewer  stameni 
than  petals. 


420 


GLOSSARY. 


Metteus.  Having  the  taste  or  smell  of 
honey. 

Melligo.    Honey-dew. 

Membranous,  Membranaceous  (-eus). 
Thin  and  rather  soft  or  pliable,  like 
a  membrane. 

Meniscoid  (-oideus).  Concavo-convex, 
like  a  meniscus. 

Mericarp  (-arpium).  One  of  the  akene- 
like  carpels  or  a  closed  half-fruit  of 
Umbelliferse ;  297. 

Merismatlc.  Dividing  into  parts  or  sim- 
ilar portions. 

Merithallus.    Synonym  of  Internode. 

Merous  (-us).  In  Greek  compounds,  de- 
notes parts  or  members ;  hence  Dime- 
rous, of  two  parts,  &c. 

Mesocarp  (-arpium).  The  middle  layer 
of  a  pericarp ;  285. 

Mesophlceum.  The  middle  or  green 
bark;  76. 

Metamorphosis,  Metamorphy,  167. 

Micropyle  (-yla).  The  spot  or  point  in 
the  seed  at  which  was  the  orifice  of  the 
orule;  277,305. 

Microspore.  The  smaller  kind  of  spore 
in  Lycopodiaceae,  &c. 

Midrib.     The  central  or  main  rib ;  92. 

Miniate  (-atus).    Vermilion-color. 

Mistus  or  Mixtus.     A  cross-breed ;  321. 

Mitrceform  or  Mitriform  (-ormis).  Mi- 
tre-shaped or  cap-shaped. 

Monadelphia.  The  Linnaean  class  con- 
taining flowers  with  Monadelphous 
stamens,  i.  e.  those  united  by  their 
filaments  into  a  tube  or  column ;  250, 
335. 

Monandria.  The  Linnaean  class  (334) 
containing  flowers  with  Monandrous, 
that  is,  a  single  stamen;  249. 

Mondnthous  (-us).    One-flowered. 

Moniliform  (-ormis).  Necklace-shaped; 
cylindrical  and  with  contractions  at 
intervals. 

Monocarpettary.     Of  one  carpel ;  261. 

Monocdrpic  (-icus).  Monocarpous,  Mon- 
ocarpian.  Only  once  fruiting;  33. 

Monocephalous  (-us).  Bearing  a  single 
capitulum. 

Monochasium.  A  cyme  with  one  main 
axis ;  152,  155. 

Monochlamydeous  (-eus).  Having  but 
one  kind  of  perianth ;  190,  340. 

Monoclinous  (-us).  Synonym  of  Herma- 
phrodite. 

Monocotyledon,  Monocotyledones,  adj. 

Monocotyledonous  (-eus).  Plants  or  em- 
bryo with  a  single  cotyledon;  23,  27, 
314,  339. 


Monolocular  (-aris).    One-celled. 
Moncecia.    Name  of  Linnaean  class  (335) 

with  flowers. 
Monoecious  (-MM),   or    Monoicous   (-««). 

With  stamens  and  pistils  in  separate 

blossoms  on  the  same  plant ;  191. 
Monogamia.     Name  of  a  Linnaean  arti- 
ficial order,  in  class  Syngenesia ;  337. 
Monograph.    A  systematic  account  of 

particular  genus,  order,  or  other  group ; 

369. 

Monoyynia.     Name  of  a  Linnaean  arti- 
ficial order,   with   solitary  pistil,   or 

style;  337;  hence,  adjectively,  Mono- 

gynous;  267. 

Monomerous.    Formed  of  a  single  mem- 
ber; 176. 
Monopetdlous     (-us).       Literally     one- 

petalled ;   but    always    used    in    the 

sense  of  Gamopetalous,  which  term  is 

to  be  preferred  ;  244. 
Monophyllous  (-us).    One-leaved. 
Monopode   ( Monopodium ),  Monopodial. 

A  stem  of  a  single  and  continuous 

axis;  55. 

Monopterous  (-us).     One-winged. 
Monopyrenus.    Containing  a  single  stone 

or  nutlet. 
Monosepalous  ( -us).    Equivalent  to  Gam- 

osepalous;    but  literally  of   a  single 

sepal ;  244. 

Monospermous  (-us).    One-seeded. 
Monostichous  (-MS).    In  a  single  vertical 

rank. 

Montstylous  (-us)  .  With  a  single  style. 
Monosymmetrical.     That  which  can  be 

bisected  into  equal  halves  in  only  one 

plane;  175. 
Mondtocous    (-w«).      Bearing    progeny 

(fruiting)  only  once,  as  annuals  and 

biennials;  33. 
Monster  (Monstrum).      A  monstrosity, 

or  unnatural  development. 
Morphology,  5. 
Moschate  (-atus).    Exhaling  the  odor  of 

musk. 
Mucilaginous,    Mucilaginosus.      Slimy ; 

of  the  consistence  or  appearance  of 

mucilage. 
Mucro,    Mucronation.      A    short    and 

abrupt  small  tip  to  a  leaf,  &c. 
Mucronate  (-atus).    Tipped  with  a  mu- 

cro;  97. 
Mucronulate  (-atus).    Minutely  mucro- 

nate. 

Mule.    A  hybrfd  or  cross-breed. 
Multidpital  ( Multiceps).    Many-headed  ; 

many  shoots  or  stems  from  the  crown 

of  one  root. 


GLOSSARY. 


421 


Multifarious  (-us).      Many-ranked,   as 

leaves  in  several  vertical  ranks. 
Multlfid  (-idus).     Cleft  into  many  lobes 

or  segments. 

Multiflorous  (-us).     Many -flowered. 
Multijugate    (Multijugus).      In     many 

pairs  or  juga. 
Multilocular    (-aris).      Many-celled    or 

several-celled. 
Multiparous.     Many-bearing ;  said  of  a 

several-branched  cyme ;  152,  155. 
Multiple  fruits.     The  fructification  of  a 

flower-cluster  when  confluent  into  one 

mass;  301. 
Multiplication.    Same  as  Augmentation  ; 

179,  200. 
Multiserial  (-alis),  Multiseriate  (-atus). 

In  several  series. 
Muricate  (-atus).   Rough  with  short  and 

firm  excrescences. 

Muriculate  (-atus}.     Minutely  muricate. 
Muscariformis.     Fly-brush-shaped . 
Musciform  (-ormis).     Moss-like  in  ap- 
pearance. 
Muscology.    The  botany  of  Mosses.    But 

is  a  hybrid  word,  and  is  replaced  by 

Bryology. 

Muticous  (-us).    Pointless,  blunt,  awn- 
less. 
Mycelium.    The  filamentous  vegetative 

growth  of  a  Fungus. 
Mycology,  Mycetology.     The  botany  of 

Fungi. 
Mycropyle.     Micropyle  misspelled. 

Naked.  Wanting  some  usual  covering ; 
as  flowers  without  perianth,  ovules 
without  coats,  seeds  not  in  a  pericarp, 
buds  without  scales. 

Napiform  (-ormis).    Turnip-shaped;  31. 

Nanus.    Dwarf. 

Natant  (-ans).  Floating  or  swimming 
under  water. 

Navicular  (-aris).  Boat-shaped.  Same 
as  Cymbiform. 

Nebulose  (-osus).     Clouded  or  misty. 

Neck.     See  Collum. 

Necklace-shaped.     See  Moniliform. 

Nectar.  The  sweetish  secretion  by  va- 
rious parts  of  the  blossom  from  which 
bees  make  honey. 

Nectary  (Nectarium).  The  place  or 
thing  in  which  nectar  is  secreted : 
formerly  applied  also  to  any  anoma- 
lous part  or  appendage  of  a  flower, 
whether  known  to  secrete  honey  or 
not ;  especially  to  the  hollow  spurs  of 
a  Violet,  Larkspur,  Columbine,  and 
the  like. 


Nectariferous  (-us).    Nectar-bearing. 
Needle-shaped.    See  Acerose. 
Nemorosus,      Nemwalis.        Inhabiting 

groves. 
Nervation.    Same  as  Venation,  or  un- 

branched  venation. 

Nerve  (Nervus).  In  botany,  this  is  a 
simple  or  unbranched  vein,  or  a  slen- 
der rib. 

Nerved,  Nervose  (-osus),  Nervate  (-atus). 
•     Having     nerves     in     the     botanical 

sense. 
Nervulose  (-osus).    Diminutive  of  ner- 

vose. 
Netted.    Same  as  Reticulated ;  Netted- 

veined;  92. 
Neurose    (-osus).      Same    as   Nervose. 

Neura  being  the  Greek  for  nerve. 
Neuter,  Neutral.     Sexless  ;  as  a  flower 
which  has  neither  stamen  nor  pistil: 
191,  195. 

Niger.     Black  or  blackish. 
Niyricans.    Turning  black  or  verging 

to  black. 

Nitidus.     Smooth  and  shining. 
Nivdlis.     Growing  in  or  near  snow. 
Niveus.     Snow-white. 
Nodding.     Hanging  down. 
Node  (Nodus).    Literally  a  knot;  the 
portion    of   a  stem   which    normally 
bears  a  leaf  or  whorl  of  leaves  ;  6. 
Nodose  (-osus).     Knotty  or  knobby. 
Nodulose  (-osus).   Diminutive  of  Nodose. 
Nomenclature,  3,  345. 
Normal    (-alis).      According    to    rule; 

agreeing  with  type. 
Notate  (-atus).  Marked  by  spots  or  lines. 
Nothus.    False  or  bastard. 
Notorhizal  (-izus).     Synonym  of  incum- 
bent,  as    applied  to  the  embryo  of 
Cruciferse. 

Nuciform  (-ormis).     Nut-like  in  shape. 
Nucleus.    A  kernel  of  an  ovule,  seed, 
&c. ;  277.     A  soft  solid  interior  part 
of  a  vegetable  cell  in  the  early  condi- 
tion; 309. 

Nuculanium.  Name  given  by  Richard 
to  a  drupaceous  or  baccate  fruit  con- 
taining more  than  one  stone  or  stony 
seed;  adopted  by  Lindley  for  a  supe- 
rior stony-seeded  berry,  such  as  a 
grape. 
Nucule  (-ula).  A  diminutive  nut  or 

stone ;  same  as  Nutlet ;  296. 
Nucumentaceus  (-eus).   Nut-like  in  char- 
acter. 
Nudicaulis.  Naked-stamened  ;  stem  not 

leafy. 
Nudus.    Naked,  in  its  various  senses. 


422 


GLOSSARY. 


Numerous  (Numerosus).  Used  in  the 
sense  of  indefinite  in  number. 

Nut  (Nux).  A  hard  and  indehiscent 
one-seeded  pericarp  resulting  from  a 
compound  ovary;  295. 

Nutlet.  A  diminutive  Nut.   See  Nucule. 

Nutant  (-an*).    See  Nodding. 

Ob.  Orer  against ;  as  a  prefix  denotes 
inversely  or  oppositely,  as 

Obcomprtssed  (-us).  Flattened  the  other 
way,  antero-posteriorly  instead  of  lat- 
erally. 

Obconical  (-icus).  Conical,  but  attached 
at  the  apex. 

Obcordate  (-atus).  Inverted  heart- 
shaped,  the  notch  at  the  apex;  97. 

Obdiplostemonous  (-us),  Obdiplostemony. 
When  the  stamens  are  double  the 
number  of  the  petals,  but  the  outer 
series  opposite  the  latter;  198. 

Obimbricate.  Imbricated  or  successively 
overlapping  downward. 

Obldnceolate.  Lanceolate  but  tapering 
toward  the  base  more  than  toward  the 
apex;  95. 

Oblique  (Obliquus).  Unequal-sided  or 
slanting. 

Oblong  ( Oblongus).  Considerably  longer 
than  broad  and  with  nearly  parallel 
sides;  95. 

Obdvate  (-atus).  Ovate  with  the  broader 
end  toward  the  apex;  95. 

Obovoid.     Solid  obovate. 

Obtectus.    Covered  by  something. 

Obtuse  (-usus).  Blunt  or  rounded  at  the 
extremity ;  97. 

Obtusiusculus.  Somewhat  obtuse ;  dimin- 
utive of  obtuse. 

Obvallatus.  Guarded  on  all  sides  or  sur- 
rounded as  if  walled  in. 

Obverse,  Obversely.  Same  as  the  prefixOb. 

Obvolute  (-utut).  A  modification  of  Con- 
volute; 138,  139. 

Ocellate  (-atus).  With  a  circular  patch 
or  a  ring  of  color. 

Ochraceous  (-eus).  Ochre  color ;  light 
/  yellow  with  a  tinge  of  brown. 

Ochrea,  Ocrea.  A  leggin-shaped  or 
tubular  stipule  or  rather  combined 
pair  of  stipules  ;  106. 

Ochreate,  Ocreate  (-atus).  Furnished 
with  ochrea  or  sheaths;  106. 

Ochroleucous  (-us).  Yellowish-white  or 
between  white  and  yellow. 

Octo.  Eight.  In  composition  gives  such 
terms  as  the  following. 

Octagynia.  Linnsean  artificial  order  with 
Octagynous  (eight-styled)  flowers ;  337. 


Octdmerous.  Composed  eight  parts  in 
the  circle. 

Octandria.  The  Linnsean  class  with 
Octandrous,  i.  e.  eight-stamened  flow- 
ers; 249,  334. 

Octdni.    In  eights. 

Octopetalous  (-us).   Eight-petalled. 

Ociosepalons.     With  eight  sepals. 

Octostichous  (-us).  In  eight  vertical 
ranks;  124. 

Oculate  (-atus).    Same  as  Ocellate. 

Officinal  (-alis).  Used  in  medicine  or 
the  arts,  therefore  in  the  shops. 

Offset.  A  short  lateral  shoot  for  propa- 
gation; 53. 

Oides,  Oideus,  -odes  and  -ides.  Greek 
for  likeness,  used  as  terminations,  in- 
dicate similarity  to;  as  Dianthoides, 
resembling  a  Pink. 

Oleraceous  (-eus).  Esculent  in  the  way 
of  a  pot-herb. 

Oliffos.  Greek  for  few ;  in  compounds 
giving  such  terms  as 

Oligdndrous  (-us).    With  few  stamens. 

Oliganthous  (-us).    Few-flowered. 

Oliydmerous  (-us).     Of  few  members. 

Oligospermous  (-us).    Few-seeded. 

Olivaceus  (-eus).    Olive-green. 

Omphalodium.  A  mark  (navel)  on  the 
hilumof  a  seed,  through  which  passed 
vessels  to  the  chalaza  or  rhaphe. 

One-sided.  Either  turned  to  one  side, 
or  with  parts  all  turned  one  way,  or 
unequal-sided. 

Oophoridium.  The  spore-case  for  the 
larger  spores  in  Selaginella,  &c. 

Opaque  ( Opdcus).  Mostly  used  in  the 
sense  of  not  shining  or  dull. 

Operculate  (-atus).  Furnished  with  an 
Operculum  or  lid. 

Operculum.  A  lid;  a  top' which  sepa- 
rates by  a  transverse  line  of  separation 
as  does  that  of  a  pyxis. 

Opposite  (-itus).  Set  against;  as  leaves 
over  against  each  other  when  there 
are  two  on  one  node ;  or  one  part  be- 
fore another,  as  a  stamen  before  a 
petal ;  6, 120, 178. 

Oppositifolius  (-ius).  Placed  opposite  a 
leaf,  as  is  a  tendril  or  peduncle  in 
Vitis,  &c. 

Oppositipetalous  (-us).  Placed  before  a 
petal. 

Oppositisepalous  (-us).  Situated  before 
a  sepal. 

Orbicular  (-arts),  Orbiculate  (-atus). 
Said  of  a  flat  body  with  a  circular  out' 
line;  95. 

Orchidactous,  246. 


GLOSSARY. 


423 


Order  ( Ordo),  Group  between  genus  (or 
tribe)  and  class;  328. 

Ordinal.    Relating  to  orders. 

Organdgeny  ( Or gano  genesis).  The  for- 
mation or  early  development  of  or- 
gans ;  2. 

Organography,  Organdlogy.  The  study 
of  organs  and  their  relations ;  2. 

Organs  of  Vegetation,  11. 

Orgydlis.  Six  feet  high,  or  of  the  height 
of  a  man. 

Ornithophilous.  Said  of  flowers  which 
are  habitually  fecundated  by  pollen 
brought  by  birds;  217. 

Orthoploceus.  Said  of  an  embryo  when 
incumbent  cotyledons  are  folded 
around  the  radicle,  as  in  Mustard. 

Orthos,  Greek  for  straight ;  whence  com- 
pounds such  as  the  following 

Orthostichies.     Vertical  ranks ;  121. 

Orthostichous.     Straight-ranked. 

Orthdtropous  (-M*),  Orthotropal.  De- 
notes an  ovule  or  seed  with  straight 
axis,  chalaza  at  the  insertion,  and  ori- 
fice at  the  other  end ;  277.  Has  been 
applied  to  an  embryo  with  radicle 
pointing  to  the  hilum;  312. 

Os  (oris).    The  mouth  or  orifice. 

Osseous  (-in).    Of  the  texture  of  bone. 

Ossiculus.  A  little  stone,  same  as  Pyrena. 

Ostiolate  (-atus).  Furnished  with  a  small 
orifice  or  little  door  (Ostiolum). 

Outgrowths,  209. 

Oval  (Ovalis).    Broadly  elliptical ;  95. 

Ovary  (Ovarium).  The  ovuliferous  part 
of  a  pistil;  166. 

Ovate  (Ovatus).  Of  the  shape  of  the 
longitudinal  section  of  a  hen's  egg, 
the  broader  end  basal ;  95.  Used  also 
for  an  egg-shaped  solid. 

Ovoid  ( Ovoideus).  Used  either  for  solid 
ovate  or  solid  oval,  more  properly  for 
the  latter. 

Ooulate  (-atus),  Ovuliferous.  Bearing 
ovules(. 

Ovule,  Ovulum.  The  body  in  the  flower 
which  becomes  a  seed ;  166,  276. 


Payina.     The  surface  of  any  flat  body, 

such  as  a  leaf. 
Palaceous  (-eus).   When  the  edges,  as  of 

a  leaf,  are  decurrent  on  the  support. 
Palate  (Palatum).    A  projection  in  the 

throat  of   a  personate  gamopetalous 

corolla;  248. 
Palea.  A  chaff,  or  chaff-like  bract,  such 

as  the  chaffy  scales  on  the  receptacle 

of  the  head  in  many  Composite;  also 


an  inner  bract  or  glume  in  Grasses; 
142. 

Paleaceous  {-eus).  Chaffy  ;  furnished 
with  pale«;  or  chaff-like  in  texture. 

Paleola.  A  diminutive  palea,  or  one 
of  a  secondarj'  order  ;  one  of  the 
names  of  the  Lodicule  or  Squamella 
in  Grasses. 

Paleolate  (-atus).  Furnished  with  pale- 
olae. 

Palets  (Pales  of  some  English  botanists). 
SameasPaleae;  142. 

Palmdris.  A  palm's  breadth  or  length; 
i.  e.  equalling  the  breadth  of  the  four 
fingers  of  the  palm. 

Palmate  (-atus).  Lobed  or  divided 
so  that  the  sinuses  point  to  or  reach 
the  apex  of  the  petiole  or  insertion ; 
101. 

Palmately  (Palmatim  or  Palmati-).  In 
the  palmate  manner. 

Palmately  veined,  93. 

Palmatifid  (-idus),  Palmatilobate,  Pnl- 
matisect).  Palmately  cleft,  lobed,  or 
divided. 

Palminerved.    Palmately  nerved ;  93. 

Pdludose  (-osus),  Palustrine  (Palustrtt 
or  Paluster).  Inhabiting  marshes. 

Pdndurate  (-atus),Panduriform  (-ormu* 
See  Fiddle-shaped. 

Panicle  (Panicula).  A  loose  compound 
flower-cluster,  such  as  is  produced  by 
the  branching  of  a  raceme,  or  the  ir- 
regular branching  of  a  corymb ;  150. 

Panicled,  Paniculate  (-atus).  In  a  pani- 
cled  manner  or  borne  in  a  panicle. 

Pannosus,  Panniformis.  Having  the  ap- 
pearance or  texture  of  felt  or  woollen 
cloth. 

Papery,  Papyraceous.  Having  the  text- 
ure of  paper. 

Papilionaceous  (-eus).  Butterfly-like; 
applied  to  a  peculiar  polypetalous 
corolla;  184,  246. 

Papillar  (-arts),  Papillose  (-osus),  Papil- 
late (-atus).  Bearing  or  resembling 
papillae,  minute  nipple-shaped  projec- 
tions. 

Pappiferous  (-us),  Pappose  (-ostu). 
Bearing  a  pappus. 

Pappus.  Thistle-down;  thence  applied 
to  various  hairy  tufts  on  akenes  or 
fruits ;  and  thence  to  any  production 
or  structure  which  takes  the  place  of 
the  limb  of  the  calyx  on  the  akenes  of 
Compositae;  192,  295. 

Papuliferous  (-us),  Populate  (-osus). 
Covered  with  Papulae,  or  small  pirn- 


424 


GLOSSARY. 


Paracarpium.  Unused  name  given  to 
an  abortive  pistil  or  carpel. 

Paracorolla.  A  crown  or  internal  ap- 
pendage or  deduplication  of  a  corolla. 

Parallel^nerved,  veined,  &c.  Same  as 
Nerved;  91. 

Parapetalous  (-us).  Said  of  stamens, 
&c.,  which  stand  at  each  side  of  a 
petal;  178,  201. 

Pardphysis,  pi.  Paraphyses.  Jointed 
thread-like  bodies,  of  no  known  func- 
tion, accompanying  the  archegonia  of 
Mosses. 

Parasitic  (-icus).  Growing  on  or  in  and 
living  upon  another  plant  or  even  ani- 
mal; 36. 

Parastemon.  Name  rarely  applied  to  an 
abortive  stamen  or  body  in  place  of  or 
accessory  to  a  stamen;  same  as  Sta- 
minodium. 

Parastichies.  Secondary  spirals  in  phyl- 
lotaxy;  127. 

Parenchyma.  Common  or  soft  cellular 
tissue. 

Parenchymatous.  Of  the  nature  of  or 
composed  of  parenchyma. 

Paries,  pi.  parities.  The  wall  of  any 
organ. 

Parietal  (Parietalis).  Borne  on  or  re- 
lated to  the  wall;  265. 

Paripinnate  (-atus).  Even-pinnate ;  same 
as  abruptly  pinnate ;  101. 

Parted,  Partite  (-itus).  Cleft  nearly 
but  not  quite  to  base;  98. 

Parthenogenesis,  Parthenogeny.  Pro- 
duction of  seed  without  the  interven- 
tion of  pollen ;  285. 

Partial  (Partialis).  Secondary,  as  Par- 
tial involucre  (142),  peduncle  (143), 
petiole  (105),  umbel  (150),  &c. 

Partible  (PartibUis).  At  length  sepa- 
rating or  easily  to  be  separated. 

Partition.  In  one  sense  a  separated  por- 
tion or  segment ;  in  another  and  the 
more  usual,  a  wall  or  dissepiment. 

PateUiform  (-ormis).  Disk-shaped,  cir- 
cular with  a  rim,  of  the  form  of  the 
patella  or  kneepan. 

Patent  (Patens).  Spreading  ;  either 
widely  open  or  diverging  widely  from 
an  axis. 

Patentissimus.  Superlative  of  Patens; 
extremely  spreading. 

Pdtulous  (-us)  Slightly  or  moderately 
spreading. 

Paucijlorous  (-us).     Few-flowered. 

Paucifolius.     Few-leaved. 

Pear-shaped.  Obovoid  or  obconical  with 
more  tapering  base. 


Pectinate  (-atus).  Pinnatifid  with  nar- 
row and  closely  set  segments,  like 
comb-teeth. 

Peddlis.     A  foot  long  or  high. 

Pedate  (-atus).  Palmately  divided  or 
parted  with  the  lateral  divisions  two- 
cleft  ;  resembling  a  bird's  foot. 

Pedatipartitus,  -lobatus,  -sectus,  &c.  Pe- 
dately  parted,  lobed,  divided,  &c. 

Pedicel  (-ellus).  An  ultimate  flower- 
stalk  or  its  division ;  the  support  of  a 
single  flower ;  143. 

Pedicellate  (-atus).  Pedicelled,  borne 
on  a  pedicel. 

Pediculus.  Name  sometimes  used  for 
Pedicel. 

Peduncle  (Pedunculus).  A  general 
flower-stalk,  supporting  either  a  clus- 
ter or  a  solitary  flower;  in  the  latter 
case,  the  cluster  may  be  regarded  as 
reduced  to  a  single  blossom ;  143. 

Pedunculate  (-atus).  Peduncled,  borne 
on  a  foot-stalk. 

Peloria.  An  irregular  flower  become 
regular  by  a  monstrous  development 
of  complementary  irregularities  ;  186. 

Peltate  (-atus),  Peltiform  (-ormis). 
Shield-form;  target-shaped;  a  plane 
body  attached  by  its  lower  surface 
(instead  of  margin  or  base)  to  a  stalk ; 
96,  107. 

Peltinerved  (-ius).  Radiately-nerved  or 
ribbed  all  round  the  circle. 

Pelviform  (-ormis).  Basin-shaped ;  shal- 
low cup-shaped. 

Pendent  (-ens).  Hanging  on  its  stalk  or 
support. 

Pendulus  (-us),  Pendulinus.  Hanging 
more  or  less,  as  if  from  weakness  of 
the  support. 

Penicillate  (-atus),  Penicittiform  (-ormis). 
Pencil-shaped,  the  pencil  (penicillum) 
being  a  brush  or  tuft  of  hairs. 

Pennate  (-atus).    Same  as  Pinnate. 

Penniform  (-ormis).  In  the  form  of  a 
feather  or  its  plume. 

Penninerved  (-ervius).  Same  as  pin- 
nately  nerved  or  veined ;  93. 

Penta.  Greek  for  five;  gives  compounds 
such  as 

Pentacdrpellary.  Composed  of  five  car- 
pels; 261. 

Pentachcenium.  Name  of  a  pentacarpel- 
lary  fruit  otherwise  like  a  cremocarp. 

Pentadelphous  (-us).  With  stamens  in 
five  clusters;  250. 

Pentagynia.  Linnaean  artificial  order 
characterized  by  Pentagynous,  i.  e. 
five-styled  flowers;  337. 


GLOSSARY. 


425 


Pentdmerous  (-us).  Composed  of  five 
members  in  a  circle;  176. 

Pentandria.  The  Linnaean  class  with 
Pentandrous,  i.  e.  five-stamened  flow- 
ers; 249,  334. 

Pentapetalous  (-us).    Five-petalled ;  244. 

Pentaphyllous  (-us).    Five-leaved;  243. 

Pentdpterous  (-us).    Five-winged. 

Pentasepalous  (-MS).     Of  five  sepals ;  244. 

Pentdstichous  (-us).  In  five  vertical  ranks ; 
123. 

Pepo,  Peponida,  Peponium.  A  gourd- 
fruit;  298. 

Perennial  (Perennis,  Perennans).  Last- 
ing year  after  year ;  32. 

Perfect  (Perfectus).  Said  of  a  flower 
which  is  hermaphrodite. 

Perfoliate  (-atus).  Where  a  stem  seem- 
ingly passes  through  a  leaf;  167. 

Perforate  (-atus).  Pierced,  or  having 
translucent  dots  which  look  like  holes. 

Pergameneus,  Pergamentaceus.  Parch- 
ment-like in  texture. 

Peri.  Greek  for  around;  hence  such 
compounds  as 

Perianth  (Perianthium).  The  floral  en- 
velopes or  leaves  of  the  flower,  consist- 
ing of  calyx,  corolla,  or  both  ;  164,  243. 

Pericarp  (-arpium).  The  fructified 
ovary;  286. 

Pericdrpic  (-icus).  Relating  to  the  peri- 
carp. 

Perichcetial  (-Mis),  Relating  to  the 
Perichcetium,  a  set  of  bracts  around 
the  fruit-stalk  in  Mosses. 

Periclddium.  The  sheathing  base  of  a 
leaf  when  it  expands  and  surrounds 
the  supporting  branch. 

Periclinium.  Involucre  of  the  capitu- 
lum  of  Composite ;  148. 

Periderm  (-erma  or  -ermis).  Outer  bark 
or  Epiphloeum. 

Perigone,  Perigdnium.  Svnonym  of 
Perianth;  164. 

Perigynium.  Name  of  hypogynous 
bristles,  scales,  or  a  sac,  which  sur- 
rounds the  pistil  (also  the  stamens 
when  present)  of  many  Cyperaceae. 

Perigynovs  (-us).  Literally  around  the 
ovary;  said  of  organs  which  are  ad- 
nate  to  the  perianth,  or  to  this  as  con- 
nate with  the  low  r  part  of  the  pistil ; 
182. 

Peripetalous  (^us).     Around  the  petals. 

Peripherie  (-icus).  Of  or  belonging  to  the 
circumference ;  as  of  an  embryo  coiled 
round  the  outside  of  the  albumen. 

Periphoranthium.  Synonym  of  the 
involucre  of  Composite ;  148. 


Peripterus.  Surrounded  by  a  wing  or 
thin  border. 

Perisperm  (-ermium).  The  albumen  of 
the  seed,  at  least  the  exterior  and  or- 
dinary albumen;  14,  310. 

Peristome  (Peristoma  or  Peristdmium). 
The  fringe  or  other  structure  surround- 
ing the  orifice  (stoma)  of  a  Moss. 

Peritropous  (-us)  or  Peritropal.  Said 
of  a  seed  which  is  horizontal  in  the 
pericarp;  or  of  a  radicle  pointed  to- 
ward the  sides  of  the  pericarp. 

Persistent  (-ens).  Remaining  even  on 
the  fruit,  or  over  winter ;  243. 

Personate  (-atus).  Masked,  as  when  a 
bilabiate  corolla  has  a  prominent  pal- 
ate; 248. 

Pervious  (-ius).  With  an  open  passage- 
way. 

Perfuse  (-usus).     Having  slits  or  holes. 

Perula,  pi.  Ferulae.  Scales  of  leaf-buds 
and  the  like;  40. 

Perulate  (-atus).  Furnished  with  peru- 
Ice  or  scales. 

Pes,  gen.  pedis.  A  foot.  Hence  in 
Latin  compounds  Longipes,  long- 
stalked,  Brevipes,  short-stalked,  &c. 

Petal  (Petalum).     A  corolla-leaf ;  165. 

Petaline  (-inus),  Petaloid  (-oideus). 
Petal-like,  or  relating  to  petals  ;  118. 

Petalody.  Name  for  the  metamorphosis 
of  other  organs  (such  as  stamens)  into 
petals;  174. 

Petiolar  (-aris).  Borne  on  or  relating 
to  a  petiole. 

Petiolate  (-atus),  Petioled.  Having  a 
petiole. 

Petiole  (Petiolus).  The  footstalk  of  a 
leaf;  85,  104. 

Petiolulate  (-atus),  Petiolular  (-arit). 
Having  a 

Petiolule  (Petiolulus).  A  footstalk  of  a 
leaflet;  105. 

Petroeus.    Growing  among  rocks. 

Petrosus.     Growing  in  stony  places. 

Phcenogams,  Phcenogamia,  Phcenoga- 
mous  plants.  Plants  sexually  propa- 
gating by  flowers,  of  which  the  essen- 
tial organs  are  stamens  and  pistil ;  3, 
334,  340,  344. 

Phalanges,  sing.  Phalanx.  The  bundles 
of  stamens  in  diadelphous  or  polyadel- 
phous flowers. 

Phanerogams,  Phanerogamia,  &c.  See 
Phaenogams,  &c. 

Phlceum.     Greek  name  for  bark. 

Phoeniceus.  Deep  red  with  some  scar- 
let. 

Phoranthium.     A  name  for  the  recep* 


426 


GLOSSARY. 


tacle  of  the  capitulum  in  Composite ; 
148. 

Phycology.    The  botany  of  Algae. 

Phylla.  Leaves  in  Greek;  combined 
with  Greek  numerals,  forming  such 
terms  as  Diphyttous,  Triphyllous,  &c., 
to  Polyphyllous. 

Phyttocladium.  A  branch  assuming  the 
function  of  foliage ;  65. 

Phyllodineous  (-eus).    Relating  to  a 

Phyllddium.  A  petiole  usurping  the 
form  and  function  of  a  leaf-blade; 
110. 

PhyUody,  Phyllomorphy.  Names  for 
the  transformation  or  metamorphosis 
of  floral  organs  into  leaves ;  174. 

Phyllotdxis,  Phyllotaxy.  Leaf-arrange- 
ment; 119. 

Phyllomania.  The  unusual  or  abnormal 
production  of  leaves. 

Phyllophore  (-orum).  The  budding  sum- 
mit of  a  stem  on  which  leaves  are  de- 
veloping. 

Phyllum.  Greek  for  leaf;  6,  85.  See 
Phrlla. 

Phyllome,  Phylldma.  An  assemblage 
of  leaves,  or  of  incipient  leaves  in  a 
bud.  Also  recently  used  by  German 
botanists  for  leaf  generically  or  poten- 
tially, that  which  answers  to  a  leaf;  6. 

Phytdgraphy.  Botany  as  relates  to  the  de- 
scription and  illustration  of  plants ;  345. 

Phy'ology.     Synonym  of  Botany. 

Phytomer,  pi.  Pkytomera.  Plant-ele- 
ments in  morphology  ;  same  as 

Phyton.  Greek  name  for  plant;  has 
been  used  in  the  sense  of  plant-ele- 
ment, or  plant-unit ;  7. 

Phytotomy.  Same  as  Vegetable  Anatomy 
or  Histology ;  2. 

Piceus.     Pitch-black  or  brownish-black. 

Pictus.    Painted,  or  rather  as  if  painted. 

Pileate  (-atus),  Pileiformis.  Having  the 
form  of  a  cap  or  Pileus. 

Pileorhiza.     The  root-cap. 

Piliferous  (-us).  Bearing  or  tipped  with 
hairs  (pill). 

Pilosciusculus.     Slightly  hairy. 

Pilose  (-osus).  Hairy,  in  general  with 
any  sort  of  pilosity ;  in  particular  with 
soft  and  distinct  hairs. 

Pinna.  One  of  the  primary  divisions  of 
a  pinnate  leaf,  either  simply  pinnate, 
when  it  is  a  leaflet,  or  a  partial  petiole 
or  rhachis  with  the  leaflets  when  the 
leaf  is  bipinnate ;  104. 

Pinnate  (-atus).  When  leaflets  are 
arranged  along  each  side  of  a  com- 
mon petiole ;  100. 


Pinnately  cleft,  lobed,  parted,  &c.;  99. 
Pinnately  veined.      Feather-veined;  93. 
Pinndtifid  (-idus).     Pinnately  cleft    v  . 
Pinnatilobatus,  Pinnatilobus.    Pinnately 

lobed. 

Pinnatipartitus.     Pinnately  parted. 
Pinnatisectus.     Pinnately  divided  quite 

down  to  the  rhachis. 
Pinnule  (Pinnula).   One  of  the  pinnately 
disposed  divisions  of  a  pinna ;  a  sec- 
ondary pinna;  104. 

Pisiform  (-ormis).  Pea-shaped;  resem- 
bling a  pea. 

Pistil  (Pistillum).  The  female  organ  of 
a  flower,  consisting  of  ovary,  style, 
and  stigma,  or  at  least  of  ovary  and 
stigma ;  302,  259. 

Pistillate  (-atus),  Pistilliferous.     Said  of 

a  plant  or  a  blossom  provided  with 

pistil,  most  properly  for  one  having 

pistil  only;  191. 

Pistillidium.     One  of  the  names  of  the 

analogue  of  pistil  in  Mosses,  &c. 
Pistillody.    Name  for  the  metamorphosis 

of  other  organs  into  carpels ;  174. 
Pitcher.     See  Ascidium.     A  tubular  or 
cup-shaped  leaf,  which  usually  holds 
some  liquid;  111. 

Pith.    A  central  cellular  part  of  a  stem, 

especially  of  an  exogenous  stem ;  75. 

Pitted.     Marked  with  small  depressions 

or  pits. 

Placenta.  That  in  the  ovary  which 
bears  the  ovules,  sometimes  the  mere 
united  margins  of  the  carpel-leaves, 
sometimes  a  thickening  or  enlarge- 
ment of  them,  or  even  of  some  other 
part  of  the  ovary ;  261. 
Placentation  (-to).  The  disposition  of 

the  placentae. 
Placentiform    (-ormis).      Quoit-shaped, 

or  in  form  like  a  flat  cake. 
Plaited.     See  Plicate. 
Plane  (Planus).    With  flat  surface  or 

surfaces. 

Platys.  Greek  for  wide,  in  such  com- 
pounds as  Platyphyllus,  broad-leaved, 
&c. 

Pleivs.     Greek   for  full,   used  in   com- 
pounds for  several  or  many ;  as  Pleio- 
phyllovit,    several-leaved,    &c.     Simi- 
larly Pleistos  for  a  great  many. 
Pleiochdsium.     A  several-rayed  cvme} 

152,  155. 

Plenus.     Full.     Flos  plenus  is  what  gar- 
deners call  a  "double  flower,"  that  is 
one  in  which  the  petals  or  other  flower- 
leaves  are  abnormally  multiplied. 
Pleurenchyma.     Same  as  woody  tissue. 


GLOSSARY. 


427 


Pleurorhizal  {-««).  Embryo  with  radicle 
against  one  edge  of  the  cotyledons; 
i.  e.  the  latter  accumbent. 

Plicate  (-atus),  Plicativus.  Folded  into 
plaits  (plicce),  usually  lengthwise ;  133, 
139. 

Plumbeus.  Lead-colored;  dull  gray 
with  some  metallic  lustre. 

Plumose  (-osus).  Feathered;  when  bris- 
tles, &c.,  have  fine  hairs  on  each  side 
like  the  plume  of  a  feather,  as  the  pap- 
pus of  Thistles. 

Plumule  (Plumula).  The  bud  or  grow- 
ing point  of  the  embryo  above  the 
cotyledons;  17. 

Plures.  Many  or  several;  used  as  a 
prefix  in  Latin  words,  such  as  Pluri- 
fiorous  (-as),  several-flowered ;  Pluri- 
locular  (-am),  several-celled ;  Pluri- 
foliolate,  with  several  leaflets,  &c. , 
Plurijugate,  in  several  pairs,  &c. 

Poculiform  (-ormis).  In  the  shape  of  a 
drinking-cup  or  goblet. 

Pod.  A  dry  and  several-seeded  dehis- 
cent fruit;  strictly  a  Legume  or  a 
Silique;  288,292. 

Podium,  Podus.  A  footstalk,  stipe,  or 
other  such  support ;  used  only  in  Greek 
compounds,  as  Podocephalus,  head 
pedunculate;  Podocarpus,  fruit  stipi- 
tate ;  or  as  a  suffix,  in  such  words  as 
Leptdpodus,  slender-stalked;  Brachy- 
podus,  short-stalked,  &c. 

Podetium.    Any  stalk-like  elevation. 

Podogynium.    Same  as  Gynophore. 

Podosperm  (-ermium).  The  stalk  of  a 
seed;  276,305. 

Pogon.  Greek  for  a  beard  ;  enters  into 
various  compound  words. 

Polembryony.     See  Polyembryocy. 

Politus.  Polished ;  applied  to  a  smooth 
and  shining  surface. 

Pointless.     Same  as  Muticous. 

Pointletted.  Minutely  pointed ;  same  as 
apiculate  or  as  minutely  acuminate. 

Pollen,  Pollen-grains.  The  fecundating 
grains  or  cells  contained  in  the  anther; 
165,  256. 

Pollen-tube.  The  slender  tube  which 
begins  as  a  protrusion  of  the  inner 
coat  of  a  pollen-grain,  and  elongates 
by  growth,  at  least  when  in  contact 
with  the  stigma ;  258. 

Pollicdris.  An  inch  long;  the  length 
of  the  terminal  joint  of  the  thumb, 
pollex. 

Poliniferous  (-us).     Pollen-bearing. 

Pollinium.  A  mass  of  pollen-grains 
more  or  less  coherent ;  257,  230. 


Pollinated  (-atus).  Said  of  a  stigma 
when  supplied  with  pollen.  . 

Poly.  In  Greek  compounds,  denotes 
numerous ;  as  in 

Polyadelphia.  Name  of  a  Linnaean  ar- 
tificial order  with  stamens  Polyadel- 
phous, or  in  several  phalanges  or 
brotherhoods;  250,  335. 

Polyandria.  Name  of  a  Linnaean  class 
with  flowers  Polyandrous,  or  having, 
an  indefinite  number  of  stamens;  249, 
334. 

Polydnthous  (-us).  Many-flowered;  hi 
the  Latin  form  same  as  multiflorous. 

Polycdrpellary.     Of  many  carpels ;  261. 

Polycdrpic  (-icus).  Fruiting  many  times 
or  indefinitely;  DeCandolle's  name 
for  a  perennial  herb ;  33. 

Polycephalous  (-alus).  Consisting  of  or 
bearing  many  heads,  capitula. 

Polycoccus.    Of  several  cocci. 

Polycotyledonous  (-eus  or  es).  Having 
several  cotyledons ;  22,  314. 

Polygdmia.  Name  of  a  Linnaean  class 
having  Polygamous  flowers,  i.  e.  some 
hermaphrodite,  some  unisexual ;  191, 
335.  Also  of  Linnaean  orders  of  Syn- 
genesia;  337. 

Polygynia.  Name  of  a  Linnaean  artifi- 
cial order  with  flowers  Polygynous, 
i.  e.  containing  numerous  carpels; 
261,  337. 

Polymer ous  (-us).  Of  numerous  mem- 
bers to  each  series  or  circle. 

Polymorphous  (-us).  Of  several  or  vari- 
ous forms. 

Polypetalous  (-us).  Having  separate 
petals;  244. 

Pdlyphore  (-6rium).  A  torus  which 
bears  many  pistils,  as  that  of  a  straw- 
berry or  raspberry. 

Polyphyllous  (-us).    Many-leaved;  244. 

Polysepalous  (-us).  Of  separate  sepals; 
244. 

Polyspermous  (-MS).    Many-seeded. 

Polystemonous  (-us).  With  many  sta- 
mens. 

Polystdchyus.    Bearing  many  spikes. 

Polystylous  (-us).    Bearing  many  styles. 

Polysymmetrical.  That  which  can  be 
divided  into  similar  halves  in  several 
or  more  than  one  plane ;  175. 

Polytocous  (-us).  Bearing  progeny  (fruit- 
ing) many  times,  i.  e.  year  after  year; 
33. 

Pome  (Pomum).  Kind  of  fruit  of  which 
the  apple  is  the  type ;  298. 

Pomeridianus.    In  the  afternoon. 

Pomiferous  (-u»).    Pome-bearing. 


428 


GLOSSARY. 


Pomology.     A  treatise  on  or  the  subject 

of  fruits  considered  as  esculent. 
Porose   (-osus),   Porous.     Pierced  with 

small  holes  or  pores. 
Posterior.     In  an  axillary  flower  is  the 

side  next  the    axis  of  inflorescence; 

160. 
Posticous  (-us).     On  the  posterior  side, 

which  in  a  flower  is  that  next  the  axis 

of  inflorescence :  an  adnate  anther  is 

posticous  when  on  the  outer  side  of 

the  filament,  i.  e.  when  it  faces  the 

petals;  253. 
Pouch.    See  Silicle. 

Prcecox.  Appearing  or  developing  early. 
Prcefloration.  Same  as  ^Estivation ;  132. 
Prcefoliation.  Same  as  Vernation  ;  132. 
Prcemorse  (-orsus).  With  end  as  it  were 

bitten  off. 

Prasinus.    Grass  green. 
Pratensis.    Growing  in  meadows. 
Prickly.     Armed    with    Prickles  (56), 

which  are  outgrowths  of  the  bark  or 

rind. 

Primine.  Outer  coat  of  the  ovule ;  277. 
Primordial  (-ialis).  The  first  in  order 

of  appearance.     Primordial  leaves  are 

those  of  the  plumule. 
Prismatic  (-icus).      Prism-shaped,  with 

flat  faces  separated  by  angles. 
Procerus.    Very  tall. 
Process    (Processus).     Any    projecting 

appendage. 
Procumbent   (-ens).    Lying    along  the 

ground ;  53. 
Productus.     Produced,  i.  e.  extended  or 

prolonged  into. 
Pro-embryo,  284. 
Proles.  Progeny ;  sometimes  used  for 

race;  320. 

Proliferous  (Prolifer,  Proliferus).  Bear- 
ing progeny,  in  the  way  of  offshoots. 
Proliferation  or 
Prolification  is  usually  taken    as    the 

production  by  one  organ  of  something 

different,  such  as  the  development  of 

buds  and  plantlets  on  leaves,  of  leafy 

shoots  in  place  of  flowers,  &c. ;  73. 
Proligerous  (-us).     Same  as  Proliferous. 
Prone  (Pronus).     Lying  flat,  especially 

face  downward. 
Propdculum,  Propdgulum.  Name  of  a 

shoot,  such  as  a  runner  or  sucker  which 

may  serve  for  propagation. 
Propdffines.     Same  as  Bulblets. 
Prophylla.     Primary  leaves,  as  the  first 

leares  of  a  branch  or  axis. 
Prosenchyma.   Plant-tissue  consisting  of 

lengthened,  tubular,  or  fusiform  cells. 


Prostrate  (-atus).    Lying  quite  flat  on 

the  ground ;  53. 

Protos.  Greek  for  first;  used  in  various 
compounds,  such  as 

Protandrous,   Protandry.      See  Proter- 

androus. 

Proterdndrous,  also  Protandrous,  Pro- 
terandry.  When  the  anthers  of  a 
flower  are  in  anthesis  earlier  than  the 
stigma;  219,  220. 

Proteranthous  (-us).  Where  flowering 
precedes  leafing. 

Proterogynous,  Proterdgyny,  or  Pro- 
toyynous,  Protogyny.  When  the 
stigma  is  ready  for  its  functions  ear- 
lier than  the  anthers  of  the  same 
blossom;  219. 

Protophytes,  Protophyta.  Alga,  &c., 
the  supposed  first  plants. 

Protoplasm,  Protopldsma.  The  forma- 
tive organic  material  of  plants  and 
animals,  in  its  living  state. 

Pruinate  (-atus),  Pruinose  (-osus).  As  if 
frosted  over  with  a  bloom  or  powder. 

Pseudos.  Greek  for  false,  a  prefix  in  . 
various  compounds,  as  Pseudo-mono- 
cotyledonous ;  26. 

Pseudo-bulb.  A  thickened  and  bulb- 
like  internode  in  epiphytal  orchids ;  a 
corm. 

Pseudocarp  (-arpium).  The  principal 
or  accessory  part  of  an  anthocarpous 
fruit;  300. 

Pseudo-costate.  False-ribbed,  as  where 
a  marginal  or  intramarginal  vein  or 
rib  is  formed  by  the  confluence  of  the 
true  veins. 

Pseudospermium.  Name  given  to  any 
kind  of  one-seeded  fruit  which  is  inde- 
hiscent  and  resembles  a  seed,  such  as 
an  akene,  &c. 

Psilos.  Greek  for  naked  or  bare;  as  in 
Psilostachyus,  with  naked  spike. 

Pteridium,  Pterodium.  Names  for  the 
Key-fruit  or  Samara. 

Pteris.  Used  for  wing  in  Greek  com- 
pounds, also  for  a  Fern. 

Pteridographia.     The  botany  of  Ferns. 

Pter acarpous  (-us).    Wing-fruited. 

Pteropodus.  Wing-footed,  i.  e.  petiole 
wing-margined,  &c. 

Ptyxls.  Greek  name  for  folding,  as  of 
leaves  in  a  bud :  132,  133. 

Pubens,  Pubes.     Used  for  Pubescent. 

Pubrrulus.     Minutely  pubescent. 

Pubes.    Pubescence,  hairiness. 

Pubescent  (-ens).  Clothed  or  furnished 
with  hairs  or  down,  especially  with  soft 
or  downy  and  short  hairs. 


GLOSSARY. 


429 


Pugioniform  (-ormis).    Dagger-shaped. 

Pullus.  Dark-colored ;  dusky-brown  or 
blackish. 

Pulvereus,  Pulverulentus.  Powdered ;  as 
if  dusted  with  powdery  matter  or 
minute  grains. 

Pulvinate  (-atus),  Pulviniform  (-ormis). 
Cushion-shaped. 

Pulvinus.  A  cushion ;  name  given  to  an 
enlargement  or  swelling  close  under 
the  insertion  of  a  leaf,  or  sometimes 
to  the  swollen  base  of  a  petiole. 

Pumilus.    Low  or  little. 

Punctate  (-atus).  Dotted,  either  with 
depressions  like  punctures,  or  trans- 
lucent internal  glands,  or  with  colored 
dots. 

Puncticulate  (-atus).    Minutely  punctate. 

Pungent  (-ens).  Terminating  in  a  rigid 
and  sharp  point  or  acumination,  like  a 
prickle. 

Puniceous  (-eus).     Bright  carmine-red. 

Purpureus.  Originally  the  red  of  arte- 
rial blood;  but  our  purple  is  some- 
what dull  red  with  a  dash  of  blue  or 
violet. 

Purpurdscens.     Purplish. 

Pusillus.  Very  small,  or  weak  and  slen- 
der. 

Pustular,  Pustulate  (-atus),  Pustulose 
(-osus).  Having  low  elevations,  like 
blisters. 

Putdmen.  The  shell  of  a  nut;  the  endo- 
carp  of  a  stone-fruit ;  288. 

Pycnvs,  Greek  for  thick ;  whence  Pycno- 
cephalus,  thick-headed,  &c. 

Pygmceus.     Dwarf,  pygmy. 

Pyramidal  (-alis).     Pyramid-shaped. 

Pyrene  (Pyrena).  Same  as  Nucule  or 
Nutlet;  one  of  the  small  stones  of  a 
drupaceous  fruit ;  298. 

Pyrenarium,  Pyridium.  A  pear  or  pear- 
like  fruit,  same  as  Pomum. 

Pyrenarius.  Name  of  a  drupaceous 
pome,  as  of  Medlar  and  Crataegus. 

Pyridion.     Synonym  of  Pome. 

Pyrenocarp  (-arpium).  A  general  name 
for  any  drupaceous  fruit ;  292. 

Pyriform  (-ormis).     See  Pear-shaped. 

Pyxidute  (-atus).     Furnished  with  a  lid. 

Pi/xidium,  Pyxis.  A  capsule  with  trans- 
verse dehiscence,  making  a  lid  of  the 
upper  portion ;  293. 


Quadri-.  In  Latin  compounds,  denotes 
four;  as  Quadrangular,  Quadrifari- 
ous  (in  four  vertical  ranks),  Quadryu- 
ffate  (in  four  pairs),  &c. 


Quaternary,  Quaternate.  In  fours  or 
composed  of  four;  176. 

Quini,  Quinary  (-ius),  Quinate  (-atut). 
In  fives ;  176. 

Quinque.  Five.  In  Latin  compounds, 
giving  rise  to  such  terms  as 

Quincuncial,  in  a  Quincunx;  also  five- 
ranked;  123,  136. 

Quinquefarious  (-ius).  In  five  vertical 
ranks. 

Quinquefoliate      (-atus).       Five-leaved. 

Quinquefoliolate,  with  five  leaflets. 

Quintuple.  Dividing  into  five  parts,  or 
five-fold. 

Quintuplinerved  or  -veined.  With  mid- 
rib of  leaf  dividing  into  five  (i.  e.  two 
lateral  pairs)  above  the  base;  93. 


Race.  A  variety  of  such  fixity  that  it 
is  reproduced  by  seed;  also  used  in  a 
looser  and  more  extended  sense  for  a 
series  of  related  individuals  without 
particular  regard  to  rank  ;  320. 

Raceme  (Race'mus).  An  indeterminate 
or  centripetal  form  of  inflorescence 
with  lengthened  axis  and  equal-pedi- 
celled  flowers;  146. 

Racemiferous.     Bearing  racemes. 

Racemiform  (-ormis).  In  the  form  of  a 
raceme. 

Racemose  (-osus).  Having  the  character 
or  appearance  of  a  raceme,  or  in  ra- 
cemes.  ' 

Rachi 

Radial.     Belonging  to  the  ray. 

Radiate  (-atus).  Spreading  from  or 
arranged  around  a  common  centre, 
or  around  the  circumference  of  a  cir- 
cle ;  bearing  rays  or  ray-flowers. 

Radiately  veined.  Same  as  Palmately 
veined;  93. 

Radiatiform  (-ormis).  Said  of  a  capitu- 
lum  of  flowers  which  is  radiate  by  en- 
largement of  some  of  the  outer  flowers, 
which  however  are  not  truly  ligulate, 
as  in  species  of  Centaurea. 

Radical  (-alis).  Belonging  to  or  pro- 
ceeding from  the  root,  or  from  a  root- 
like  portion  of  stem  at  or  below  the 
surface  of  the  soil. 

Radicant  (Radicans).     Rooting. 

Radicel.     A  minute  root  or  a  rootlet. 

Radiciflorous  (-us).  Flowering  (appar- 
ently) from  the  root. 

Radiciform  (-ormis),  Radicinus.  Of  the 
nature  or  appearance  of  a  root. 

Radicle  (-icula).  Literally  a  diminutive 
root;  but  the  "  radicle  "  of  the  embryo, 


, 

es.  nJk',^**!      IQ\  I    4; 

is.    See  Rhachis.    f  •    ' 


430 


GLOSSARY. 


so  called  in  descriptive  botany,  is  the 
hypocotyledonary  and  primal  inter- 
node.  (See  Caulicle);  10. 

Radiculose  (-osus).     Bearing  rootlets. 

Radix.     The  root. 

Rameal  (-alis),  Rameus.  Belonging  to 
(Ramus)  a  branch. 

Ramenta.  Thin  chaffy  scales  belonging 
to  the  surface  or  epidermis,  such  as 
the  chaff  on  the  stalks  of  many  Ferns. 

Ramification.     Branching;  47. 

Ramiflorous  (-us).  Flowering  on  the 
branches. 

Ramose  (-osus).     Branching  or  branchy. 

Ramulose  ( -  osus ).  Bearing  many  branch- 
lets,  i.  e.  Ramuli  or 

Raphe.    See  Rhaphe. 

Rdphides  or  Rhaphides.  Crystals  in 
the  cells  of  plants,  especially  needle- 
shaped  crystals. 

Ray  (Radius).  One  of  the  radiating 
branches  of  an  umbel  (147);  also  the 
marginal  as  opposed  to  the  central  part 
(or  disk)  of  a  head,  umbel,  or  other 
flower-cluster,  when  there  is  a  differ- 
ence of  structure.  Also  used  as  an 
abbreviated  expression  for 

Ray-flowers.  Those  which  belong  to 
the  margin  of  a  circular  flower-clus- 
ter, and  differ  from  (being  usually 
larger  than)  those  of  the  disk. 

Recaulescence.  The  adhesion  of  leaves 
or  their  stalks  to  a  stem ;  158. 

Receptacle  (Receptdculum).  A  portion 
of  axis  forming  a  common  support  or 
bed  on  which  a  cluster  of  organs  is 
borne.  The  receptacle  of  the  flower, 
or  the  torus,  is  the  axile  portion  of  a 
blossom,  that  which  bears  sepals, 
petals,  stamens,  and  pistils ;  167,  211. 
The  receptacle  of  inflorescence  is  the 
axis  or  rhachis  of  the  head,  spike,  or 
other  dense  cluster ;  143. 

Reclinate  (-atus),  Reclined,  Reclining. 
Falling  or  turned  toward  downward, 
so  that  its  upper  part  rests  on  the 
ground  or  other  object;  53,  133. 

Rectinervius.  Straight-veined  or  straight- 
nerved;  92. 

Rectiserial  (-ialis).  Tn  rectilinear  ranks ; 
124. 

Recurved  (-us),  Recurvatus.  Curved 
backward  or  downward. 

Reduplicate  (-atus)  or  Reduplicativus. 
Folded  and  projecting  outward. 

Reflexed  (-us).  Abruptly  bent  or  turned 
downward  or  backward. 

Refracted  (-us).  Same  as  reflexed,  but 
abruptly  bent  from  the  base. 


Regma.  A  two-several-lobed  two- 
several-celled  fruit  (2-pluricoocous), 
which  separates  at  maturity  into  as 
many  2-valved  carpels,  as  in  Euphor- 
bia ;  one  form  of  Schizocarp. 

Regmacarp  (-arpium).  A  general  name 
of  a  dry  and  dehiscent  fruit ,  292. 

Regular  (-am).  Uniform  in  shape  or 
structure;  symmetrical  as  respects 
shape;  175. 

Reniform  (-ormis).  Kidney-shaped; 
having  the  outline  of  the  longitudinal 
section  of  a  kidney ;  96. 

Repand  (Repandus).  With  slightly  un- 
even margin,  which,  if  more  pro- 
nounced, would  be  sinuate ;  98. 

Repent  (Repens).  Creeping,  i.  e.  pros- 
trate or  horizonal  and  rooting ;  53. 

Replicate  (-atus),  Replicativus.  Folded 
backward. 

Replum.  A  frame-like  placenta  (like 
a  door-case),  from  which  the  valves 
of  a  capsule  or  other  dehiscent  fruit 
fall  away  in  dehiscence,  as  in  Cruci- 
ferse,  certain  Papaveracese,  Mimosa, 
&c.;  293. 

Reptant  (Reptans).    Same  as  Repent. 

Resupinate.  Upside  down,  or  having 
that  appearance. 

Rete.    Network. 

Reticulated  (-atus),  Retiformis.  In  the 
form  of  network ;  netted. 

Reticulate-veined,  92. 

Retindculum.  Name  sometimes  applied 
to  the  gland  to  which  one  or  more 
pollinia  are  attached  in  Orchids,  &c. 
The  persistent  and  indurated  hook- 
like  funiculus  of  the  seeds  in  most 
Acanthaceae. 

Retinerved  (-ius).  Same  as  Reticulate- 
veined. 

Retrocurved  (-us).    Same  as  Recurved. 

Retroflexed  (-us).     Same  as  Reflexed. 

Retrorse  (-orsus).  Directed  backward  or 
downward. 

Retroverted  (Retroversus).    Inverted. 

Refuse  (Retusus).  With  a  shallow  or 
obscure  notch  at  a  rounded  apex ;  97. 

Reversion.  A  changing  back,  or  in  the 
reverse  direction ;  171. 

Revolute  (-utus).  Rolled  backward  from 
the  margins  or  apex;  133. 

Rhachis.  The  axis  (backbone)  of  a 
spike  or  of  a  compound  leaf;  101, 143. 

Rhaphe.  The  adnate  cord  or  ridge 
which  in  an  anatropous  ovule  con- 
nects the  hilum  with  the  chalaza; 
279,  307. 

Rhipidium.    A  fan-shaped  cyme ;  156. 


GLOSSARY. 


431 


Rhizanthout  (-us).  Root-flowered ;  flower- 
ing from  the  root  or  seeming  root. 

Rhizina.  The  peculiar  roots  or  root- 
hairs  of  Mosses,  Lichenes,  &c. 

Rhizocarpous  (-MS).  Rhizocarpic  (-icus). 
Literally  root-fruited;  used  by  De- 
Candolle  for  a  perennial  herb. 

Rhizome,  Rhizoma.  A  rootstock;  a 
stem  of  root-like  appearance  pros- 
trate on  or  underground,  from  which 
rootlets  are  sent  off;  the  apex  pro- 
gressively sending  up  herbaceous 
stems  or  flowering  stalks  and  often 
leaves;  56. 

Rhizomorphous  (-us).  Root-like  in  ap- 
pearance. 

Rhombic  (-icus).    Rhomb-shaped. 

Rhomboidal  (-alls).  Approaching  a 
rhombic  outline;  quadrangular,  with 
the  lateral  angles  obtuse. 

Rib.  A  primary  and  strong  vein  or 
conspicuous  portion  of  the  framework 
of  a  leaf;  92. 

Ribbed.  Furnished  with  prominent 
ribs. 

Rictus.  The  mouth  or  gorge  of  a  bila- 
biate corolla. 

Rima.    A  chink  or  cleft. 

Rimose  (-osus).  With  chinks  or  cracks, 
like  those  of  old  bark. 

Ring.    In  Ferns,  &c.    See  Annulus. 

Ringent  (-ens).  Grinning  or  gaping  ;  as 
is  the  mouth  of  an  open  bilabiate 
corolla;  248. 

Riparita.  Growing  along  the  banks  of 
rivers,  &c. 

Rivdlis.    Growing  along  brooks. 

Rivularis.  Growing  in  watercourses  or 
rivulets. 

Root.  The  descending  axis.  Roots  are 
axes  which  grow  in  the  opposite  di- 
rection from  the  stem,  are  not  com- 
posed of  nodes  and  internodes,  are 
mostly  developed  underground,  and 
absorb  moisture,  &c.,  from  the  soil; 
27. 

Root-cap,  13,  28. 

Root-hairs.  Attenuated  unicellular 
outgrowths  or  hairs  from  the  newly 
formed  parts  of  a  root,  for  absorp- 
tion; 13,  29. 

Rootlet.  A  very  slender  root  or  branch 
of  a  root. 

Rootstock.     See  Rhizoma ;  56. 

Roridus.  Dewy ;  covered  with  particles 
resembling  drops  of  dew. 

Rosaceous  (-eus).  Arranged  like  the 
five  petals  of  a  normal  rose;  246. 
Sometimes  used  for  rose-color. 


Roseus.    Rose-colored ;  pale  red. 

Rostellate  (-atus).  Diminutive  of  Ros- 
trate. 

Rostellum.  A  diminutive  beak.  Also 
the  name  applied  by  Linnaeus  to  the 
Caulicle  or  Radicle. 

Rostrate  (-atus).  With  a  Rostrum,  a 
beak  or  spur ;  narrowed  into  a  slender 
tip  or  process. 

Rosular,  Rosulate  (-atus).  Collected  in 
a  rosette. 

Rotate  (-atus).  Wheel-shaped;  circular 
and  horizontally  spreading  very  flat ; 
248. 

Rotund  (Rotundus,  Rotundatus).  Round- 
ed in  outline;  95. 

Rough,  Roughish.     See  Scabrous. 

Rubellus,  Rubescent  (-ens),  Rubens.  Red- 
dish. Rubescent  also  is  turning  red. 

Ruber.     Red  in  general. 

Rubicundus.  Blushing,  turning  rosy- 
red. 

Rubiffinose  (osus).    Brownish  rusty-red. 

Ruderal  (-alis).  Growing  in  waste  places 
or  among  rubbish. 

Rudiment.  An  imperfectly  developed 
and  functionally  useless  organ;  a 
Vestige. 

Rufous  (-us),  Rufescent  (-ens).  Pale 
red  mixed  with  brown. 

Rugose  (-osus).  Covered  or  thrown  into 
wrinkles,  Rugce. 

Ruminated  (-atus).  As  if  chewed ;  said 
of  the  albumen  of  a  nutmeg,  &c. ;  311. 

Runcinate  (-atus).  Saw-toothed,  or 
sharply  incised,  the  teeth  or  incisions 
retrorse. 

Runner.  A  prostrate  filiform  branch 
which  is  disposed  to  root  at  the  end 
or  elsewhere ;  53. 

Running.    Same  as  Repent. 

Rupestris,  Rupicola.  Growing  on  rocks 
or  in  rocky  places. 

Ruptilis.     Bursting  irregularly. 

Rusty.  Same  as  Rubiginose,  Rufescent, 
and  Ferruginous. 

Rutilans.  Deep  red  with  a  metallic 
lustre. 


Sabulosus.     Growing  in  sandy  places. 
Saccate  (-atus),  Sacciform.    Sac-shaped; 

baggy. 
Sagittate   (-atus),  Sagittiform  (-ormis). 

Arrow-head-shaped. 
Salsuginosus.    Growing  within  reach  of 

salt  water. 
Salver-shaped.      See    Hypocraterimor- 

phous;  248. 


432 


GLOSSARY. 


Samara.  An  indehiscent  winged  fruit ; 
294. 

Samaroid.    Resembling  a  samara. 

Sap-wood.  New  wood  of  an  exogenous 
stem;  80. 

Sarcocarp  (-arpium).  The  succulent  or 
fleshy  portion  of  a  drupe ;  285.  Has 
been  proposed  also  as  a  general  name 
for  a  baccate  fruit;  292. 

Sarmentose  (-osus).  Producing  long 
and  lithe  branches  or  runners,  viz. 
Sarments  (Sarmenta). 

Sativus.    That  which  is  sown  or  planted. 

Saw-toothed.     See  Serrate. 

Saxdtilis,  Saxosus,  Saxicolus.  Living  on 
or  among  rocks. 

Scabridus,  Scabriusculus.  Roughish ; 
diminutive  of 

Scabrous  (Scaber).     Rough  to  the  touch. 

Scalariform  (-ormis).  Ladder-shaped; 
with  transverse  markings  like  the 
rounds  of  a  ladder. 

Scales.  Any  thin  scarious  bodies,  usu- 
ally degenerate  leaves,  sometimes  of 
epidermal  origin. 

Scalloped.     Same  as  Crenate ;  98. 

Scaly.     See  Scarious,  Squamose. 

Scaly  Buds,  40. 

Scandent  (-ens).  Climbing,  in  what- 
ever mode ;  51. 

/Scope  ( Scapus).  A  peduncle  rising  from 
the  ground;  51,  143. 

Scapiform  (-ormis),  Scapose  (-osus).  Re- 
sembling a  scape. 

Scapigerous  (-us).    Scape-bearing. 

Scar.  The  mark  left  on  the  stem  by 
the  separation  of  a  leaf,  or  on  a  seed, 
&c.,  by  its  detachment. 

Scarious  or  Scariose  (-osus).  Thin,  dry, 
membranaceous,  and  not  green. 

Schizocarp  (-arpium).  A  pericarp  which 
splits  into  one-seeded  pieces ;  296. 

Scion.  A  young  shoot ;  a  twig  used  for 
grafting. 

Sciuroideus.     Like  a  squirrel's  tail. 

Scleranthium.  Name  of  the  fruit  of 
Mirabilis,  and  the  like;  an  akene 
enclosed  in  an  indurated  portion  of 
calyx-tube. 

Scleroideus.  Having  a  hard  texture; 
from  Scleros,  hard. 

Scobiform  (-ormis).  Having  the  appear- 
ance of  sawdust. 

Scorpioid.  A  form  of  unilateral  inflo- 
rescence which  is  circinately  coiled  in 
the  bud :  in  the  stricter  sense,  a  form 
with  the  flowers  two-ranked,  these 
being  thrown  alternately  to  the  right 
and  left;  155,157. 


Scrobiculate  (  atus).  Marked  by  minute 
or  shallow  depressions. 

Scrotiform  (-ormis).     Pouch-shaped. 

Scurf.  Small  and  bran-like  scales  on 
the  epidermis. 

Scutate  (-atus),  Scutiform  (-ormis). 
Buckler-shaped. 

Scutelliform  (-ormis).    Platter-shaped. 

Scymetar-shaped.     See  Acinaciform. 

Sectile  (-ills).    As  if  cut  up  into  portions. 

Section  (Sectio).  In  classification,  is 
applied  in  a  general  way  to  a  divi- 
sion in  the  arrangement  of  genera, 
species,  or  other  groups ;  327. 

Sectus.     Completely  divided ;  99. 

Secund  (Secundus).  When  parts  or 
organs  are  all  directed  to  one  side. 

Secundiftorus.  With  flowers  of  a  cluster 
all  secund. 

Secundine.  The  second  (inner)  coat 
of  an  ovule ;  277. 

Seed.  The  fertilized  and  matured  ovule ; 
the  result  of  sexual  reproduction  in  a 
phsenogamous  plant;  305. 

Seed-leaves.     Cotyledons,  11. 

Seed-stalk.  See  Funiculus  and  Podo- 
sperm. 

Seed-vessel.     See  Pericarp. 

Segetdlis.     Growing  in  grain-fields. 

Segment  (Segmentum).  One  of  the 
divisions  into  which  a  plane  organ, 
such  as  a  leaf,  may  be  cleft. 

Segregate  (-atus).  Separated;  kept 
apart. 

Semen.    Seed. 

Semi.  Half,  in  Latin  compouris ;  such  as 

Semi-adherent.  The  lower  half  adhe- 
rent, &c. ;  Semi-amplexi&iul  (-aulis), 
half  clasping  the  stem  ;  Semiovate, 
ovate  halved  lengthwise,  &K. 

Semiandtropous.  Same  af  Amphitro- 
pous ;  279. 

Semilunar,  Semilunate  (-atu*).  A  syn- 
onym of  Lunate,  being  J'ke  a  half- 
moon. 

Seminal  (-alis).     Relating  tt>  the  seed. 

Seminiferous  (-us).     Seed-bearing. 

Sempervirent  (Semperviren»\.  Ever- 
green. 

Stnary  (-arrus).     In  sixes;  176. 

Sepal  (Sepalum).     A  calyx-leaf ;  165. 

Sepaline  (-inus),  Sepalous.  Relating  to 
sepals. 

Sepaloid  (-oideus).    Resembling  a  sepal. 

Sepalody.  Name  for  the  metamorphosis 
of  petals,  &c.,  into  sepals  or  seoaloid 
organs;  174. 

Separated  flowers.  Those  of  distinct 
sexes ;  same  as  Diclinous ;  191. 


GLOSSARY. 


433 


Septate  (-atus).  Separated  by  a  parti- 
tion or  septum. 

Septicide,  Septiddal  (-cidus).  When  a 
capsule  dehisces  through  the  dissepi- 
ments or  lines  of  junction  ;  289. 

Septiferous  (-us).  Bearing  the  partition 
or  dissepiment. 

Septifragal  (-us).  Where  the  valves  in 
dehiscence  break  away  from  the  dis- 
sepiments; 290. 

Septum.  Any  kind  of  partition,  whether 
a  proper  dissepiment  or  not. 

Septulate  (-atus).  Divided  by  spurious 
or  transverse  septa. 

Serial  (Serialis)  or  Seriate  (Seriatus). 
Disposed  in  series  or  rows,  whether 
transverse  or  longitudinal. 

Sericeous  (-eus).  Silky;  clothed  with 
close-pressed  soft  and  straight  pubes- 
cence. 

Serotinous  (-MS).  Produced  compara- 
tively late  in  the  season. 

Serrate  (-atus).  Beset  with  antrorse 
teeth;  97. 

Serrulate  (-atus).  Serrate  with  very 
small  or  fine  teeth;  97. 

Sesqui.  A  Latin  prefix  denoting  one 
and  a  half ;  as,  Sesquipedalis,  a  foot 
and  a  half. 

Sessile  (-ilis).  Sitting  close,  without  a 
stalk ;  destitute  of  peduncle,  pedicel, 
or  petiole,  as  the  case  may  be. 

Seta.    A  bristle,  or  bristle-shaped  body. 

Setaceous  (-eus).     Bristle-like. 

Setiform  (-ormis).  In  the  form  of  a 
bristle. 

Setigerous  (-us).    Bristle-bearing. 

Setose  (-osus).  Beset  with  or  abounding 
in  bristles ;  bristly. 

Setula.    Diminutive  of  Seta. 

Setulose  (-osus).  Bearing  or  consisting 
of  minute  bristles. 

Sea;.  Latin  for  six;  as  in  Sexangular, 
Sexfarious,  Sexpartile,  &c. 

Shaggy.  Pubescent  with  long  and  soft 
hairs ;  same  as  Villous. 

Sheath.    A  tubular  or  enrolled  part  or 
organ,  such  as  the  lower  portion  of 
the  leaf  in  Grasses.     See  Vagina. 
Sheathing.    Enclosing  as  by  a  sheath. 
Shield-shaped.   In  the  form  of  a  buckler ; 
plane  and  round  or  oval,  with  stalk 
attached  to  some  part  of  the  under 
surface;  96.     See  Clypeate,  Scutate, 
Peltate. 
Shrub.    A  woody  perennial  of  less  size 

than  a  tree ;  50. 

Shrubby.  Having  the  character  of  a  shrub. 
Sieve-cells,  77. 


Sigillate  (-atus).    As  if  marked  with  the 

impression  of  a  seal,  as  the  rootstock 

of  Polygonatum. 
Sigmoid  (-oideus).    Doubly  curved  like 

the  Greek  ?  or  the  capital  S. 
Silicle  (Silicula).    A  short  silique,  not 

very  much  longer  than  wide ;  294. 
Siliculosa.    Name  of  the  Linnsean  arti- 
ficial order  of  the  class  Tetradynamia, 

having  Siliculose  pods  ;  337. 
Silique  (Siliqua).     The  peculiar  pod  of 

Cruciferae,     especially    when    much 

longer  than  wide;  293. 
Siliquosa.    Name  of  the  other  order  of 

Tetradynamia,   with   Siliquose    fruit, 

i.  e.  a  Silique ;  337. 
Silky.  See  Sericeous. 
Silver-grain.  The  glittering  plates  in 

exogenous    wood    belonging   to    the 

medullary  rays;  74. 
Simple  (Simplex).    Of  one  piece,  series, 

&c.     A  simple  pistil  is  of  one  carpel ; 

a  simple  leaf,  of  one  blade,  &c. 
Simple  Fruits,  291. 

Simplicissimus.    Most  simple ;  complete- 
ly simple. 
Sinistrfirse.    Turned  or  directed  to  the 

left ;  51,  140. 
Sinuate  (-atus).     With  a  strongly  wavy 

or  recessed  margin ;  98. 
Sinus.    A  recess  or  re-entering  angle. 
Slothed.    Same  as  Laciniate. 
Smooth.     Either  opposed  to  scabrous, 

i.  e.  not  rough,  or  to  glabrous,  t.  ft 

not  pubescent ;  the  former  is  the  moie 

correct  application. 
Soboles.  Shoots,  especially  those  from 

the  ground. 
Soboliferous   (-MS).      Bearing  vigorous 

lithe  shoots. 

Solid  Bulb.    A  conn ;  61. 
Solitary  (-arius).   Single,  only  one  from 

the  same  place. 
Solubilis.     Separating  into  portions  or 

pieces. 

Solutus.    Loosed ;  becoming  separate. 
Sordidus.    Of  a  dull  or  dirty  hue. 
Sorediate  (-atus).   Bearing  small  patches 

on  the  surface. 
Sorfrna.   A  heap  of  carpels  belonging  to 

one  flower;  263. 
Sori,  sing,  sorus.     Heaps,  such  as  the 

clustered  fruit-dots  of  Ferns. 
Sorose.    Heaped  or  bearing  Sori. 
Sorosis.     A  fleshy  multiple  fruit,  such 
as  a  mulberry,  bread-fruit,  and  pime- 
apple. 

Spadiceus.    A  bright  and  clear  brown, 
or  chestnut  color. 


434 


GLOSSAEY. 


Spadiceous.  Having  the  nature  of  or 
bearing  a 

Spadix.   A  spike  with  a  fleshy  axis ;  149. 

Span.  The  length  of  the  space  between 
the  tip  of  the  thumb  and  that  of  the 
little  finger,  when  outstretched ;  about 
nine  inches. 

Sparsus.  Sparse  or  scattered;  whence 
Sparsiftorus,  with  scattered  flowers ; 
Sparsifolius,  with  scattered  leaves,  &c. 

Spathaceous  (-its).  Spathe-bearing,  or 
of  the  nature  of  a 

Spathe  (Spatha).  A  large  bract,  or  a 
pair  of  bracts,  enclosing  a  flower-clus- 
ter; 142. 

Spathella.  An  unused  name  for  the 
glumes  of  Grasses. 

Spathilla.  A  secondary  or  diminutive 
spat  he. 

Spatulate  (Spathulatus),  Oblong  with 
the  lower  end  attenuated,  shaped  like 
a  druggist's  spatula;  95. 

Species.  The  particular  kind,  the  unit 
in  natural  history  classification ;  317. 

Specific  Character,  Name,  &c.,  349, 
363. 

Spermaphore  or  Spermophore  (-drum). 
A  name  for  the  Placenta. 

Spermoderm  (-ermis).  The  outer  seed- 
coat;  305. 

Spermodophorum  or  Spermophorum.  An 
unused  name  for  the  gynophore  in 
Umbelliferae.  The  latter  also  an  un- 
used name  for  the  Placenta;  261. 

Spermotheca.  An  unused  name  for  peri- 
carp. 

Spermum.  Latin  form  of  the  Greek  word 
for  seed.  Lat.  Semen. 

Sphalerocarpium.  Name  proposed  for 
an  accessory  fruit,  such  as  that  of 
Shepherdia,  in  which  an  akene  is 
enclosed  in  a  baccate  calyx-tube. 

Spica.     See  Spike. 

Spicate  (-atus).  In  the  form  of  or  resem- 
bling a  spike,  or  disposed  in  spikes. 

Spiciform  (-ormis).     Spike-like. 

Spicula.  A  diminutive  or  secondary 
spike;  a  Spikelet. 

Spike  (Spica).  A  form  of  indeterminate 
inflorescence,  with  flowers  sessile  on 
an  elongated  common  axis ;  149. 

Spikelet  (Spicula).  A  secondary  spike; 
the  name  given  to  the  Locusta  or  clus- 
ter of  one  or  more  flowers  of  Grasses 
subtended  by  a  common  pair  of 
glumes. 

Spindle-shaped.     See  Fusiform. 

Spine  (Spina).  A  sharp-pointed  woody 
or  indurated  body,  commonly  a  branch, 


sometimes  a  petiole,  stipule,  or  other 
part  of  a  leaf;  55,  117. 

Spinescent  (-ens).  Ending  in  a  spine 
or  sharp  point ;  55. 

Spinose  (-osus).  Furnished  with  spines, 
or  of  a  spiny  character ;  55. 

Spinuliferous  or  Spinulose  (-osus).  Fur- 
nished with  diminutive  spines  or  Spin- 
ula. 

Spiral  (Spirdlis).  As  if  wound  round 
an  axis.  Spiral  Ducts,  68.  Spiral 
PhyUotaxy,  119,  121. 

Spiricles.  The  delicate  coiled  threads 
in  the  hairs  on  the  surface  of  certain 
seeds  and  akenes,  which  uncoil  when 
wet;  307. 

Spithamceus.  A  span  long ;  the  length 
spanned  between  the  tip  of  thumb  and 
forefinger  when  extended. 

Splendens.    Resplendent  or  glittering. 

Spongelet,  Spongiole  (-iola).  Name 
given  to  young  root-tips;  once  sup- 
posed to  be  a  peculiar  organ ;  28. 

Sporadic  (-icus).  Widely  dispersed  or 
scattered. 

Sporangium.  A  spore-case  or  theca  con- 
taining the  analogues  of  seeds  (spores) 
in  the  higher  Cryptogams. 

Spore  (Spora,  Greek  for  seed).  The 
analogue  of  seed  in  Cryptogams. 

Spore-case.     See  Sporangium. 

Sporidium.  Synonym  or  diminutive  of 
Spore. 

Sporiferous.    Spore-bearing. 

Sp&rocarp  (-arpium).  Name  given  to 
certain  spore-cases,  as  of  Lycopodi- 
aceae. 

Sporophore  (-orum).  One  of  the  syno- 
nyms of  Placenta. 

Sporule  (Sporula).  Diminutive  spore  or 
a  sort  of  spore. 

Sporuliferous  (-us).  Bearing  or  con- 
taining spores. 

Sport.  A  bud-variation  or  seed-varia- 
tion; 319. 

Spumescent  (-ens),  Spumose.  Froth-like 
in  appearance. 

Spur.  A  hollow  and  slender  extension 
of  some  portion  of  the  blossom,  usu- 
ally nectariferous,  as  of  the  calyx  of 
Larkspur  and  the  corolla  of  Violet: 
rarely  applied  also  to  a  solid  spur-like 
process. 

Spurred.  Producing  a  spur.  See  Cal- 
carate. 

Squama.  A  scale  of  any  sort,  usually 
the  homologue  of  a  leaf. 

Squamate  (-atus),  Squamiferous,  Squa- 
mosus.  Furnished  with  scales. 


GLOSSARY. 


435 


Squamella,  Sqvdmula.  Diminutive 
squama;  scales  of  secondary  order 
or  reduced  size. 

Squamiform  (-ormis).     Scale-like. 

Squamulose  (-osus).  Covered  or  beset 
with  minute  scales. 

Squarrose  (-osus).  Literally  rough- 
scurfy;  applied  to  bodies  rough  with 
spreading  and  projecting  processes, 
such  as  tips  of  bracts,  &c. 

Squarrulose  (-osus).  Diminutively  squar- 
rose. 

Stachys.    Greek  for  spike. 

Stalk.  Any  kind  of  lengthened  support 
on  which  an  organ  is  elevated. 

Stamen.  One  of  the  elements  or  phylla 
of  the  androecium  ;  165. 

Stamineal,  Stamineous  (-eus).  Relating 
to  the  stamens ;  191. 

Staminiferous  (-MS).     Stamen-bearing. 

Staminodium.  A  sterile  stamen,  or 
what  answers  to  a  stamen,  whatever 
its  form,  without  anther. 

Staminody.  Name  for  the  metamor- 
phosis of  other  floral  organs  into 
stamens;  174. 

Standard.  The  posterior  petal  of  a 
papilionaceous  corolla;  184. 

Slant.  Supporting  itself  in  an  erect 
position. 

Station.  Particular  place  as  to  soil,  ex- 
posure, &c.,  which  a  plant  affects; 
366. 

Stellate  (~atus).  Star-shaped,  arranged 
like  the  rays  or  points  of  a  star. 

Stellulate  (-attts)  or  Stellular.  Dimin- 
utive of  Stellate. 

Stem.  The  main  ascending  axis  of  a 
plant;  45. 

Stemlesi.  See  Acaulescent ;  with  no  leaf- 
bearing  stem  above  ground ;  45. 

Stemlet.  Diminutive  stem;  as  that  of 
the  plumule. 

Stenos.    Greek  for  narrow;  hence 

Stenophyllus.     Narrow-leaved,  &c. 

Steriama.  Any  foliaceous  prolongation  of 
the  blade  of  a  leaf  down  on  the  stem 
by  decurrence. 

Sterigmum.  Name  of  Desvaux  for  the 
Dieresilis  of  Mirbel. 

Sterile  (-ills).  Barren,  as  a  blossom 
destitute  of  pistil,  191 ;  a  stamen 
without  anther,  or  an  anther  without 
pollen ;  an  ovary,  without  good  ovules, 
seeds  without  embryo,  &c.  In  com- 
mon English  use,  a  male  or  staminate 
flower  is  said  to  be  a  sterile  flower. 

Stichus.  Greek  for  row  or  rank,  usually 
meaning  vertical  rank  ;  hence  such 


compounds  as  Distichous,  two-ranked ; 
Tristichous,  three-ranked,  &c. 

Stiff  ma,  pi.  stigmata.  That  part  or  sur- 
face of  a  pistil  (usually  on  or  a  part  of 
the  style,  or  in  place  of  it)  which  re- 
ceives the  pollen  for  the  fecundation 
of  the  ovules;  166. 

Stigmatic  (-ictw),  Stigmatoie  (-osiu). 
Relating  to  stigma. 

Stigmatiferous.    Stigma-bearing. 

Stings.  Stinging  hairs,  seated  on  a 
gland  which  secretes  an  acrid  liquid, 
as  in  Nettles. 

Stipe  ( Stipes).  A  stalk  of  various  sorts ; 
the  support  of  the  cap  of  a  mush- 
room ;  the  leafstalk  of  a  Fern  ;  any 
stalk-like  support  of  a  gynoecium  or 
a  carpel;  212. 

Stipd  (Stipellum).  An  appendage  to  a 
leaflet  analogous  to  the  stipule  of  a 
leaf;  106. 

Stipellate  (-atus).  Provided  with  stipels ; 
106. 

Stipitate  (-atus).  Having  a  stipe  or 
special  stalk. 

Stipitiform  (-ormis).  Shaped  like  a  stipe  | 
stalk-like. 

Stipulaceous  (-eus),  Stipular  (-am). 
Belonging  to  stipules. 

Stipulate.    Possessing  stipules. 

Stipules.  Appendages  or  adjuncts  of  a 
leaf  one  on  each  side  of  the  insertion ; 
85,  105. 

Stirps,  pi.  stirpet.    A  race. 

Stock.  Synonym  of  Race;  also  the 
portion  of  a  stem  to  which  a  graft  is 
applied;  a  caudex,  rhizoma,  or  root- 
like  base  of  a  stem  from  which  roots 
proceed;  51. 

Stole,  Stolon  (Stolo).  A  sucker,  runner, 
or  any  basal  branch  which  is  disposed 
to  root;  53. 

Stoloniferout  (-«*).  Sending  of  or  propa- 
gating by  stolons,  runners,  &c. 

Stoma,  pi.  stdmala,  Stomate.  One  of 
the  apertures  in  the  epidermis  of  folia- 
ceous parts,  through  which  cavities 
within  communicate  with  the  external 
air;  89. 

Stomatiferous  (-««).  Bearing  stomata 
or  "breathing  pores." 

Stone.    The  hard  endocarp  of  a  drupe. 

Stone-fruit.  A  Drupe,  such  as  a  peach 
or  plum ;  297. 

Stool.  The  plant  from  which  layers  are 
propagated,  by  bending  down  to  the 
ground  to  be  rooted. 

Stramineous  (-eus).  Straw-like  or  straw- 
colored. 


436 


GLOSSARY. 


Strap-tkaped.  See  Ligulate  (247)  and 
Lorate. 

Btriate  (-atta).  Marked  with  fine  longi- 
tudinal lines,  streaks,  or  diminutive 
grooves  or  ridges  (Stria). 

Strict  (Strictus).  Close  or  narrow  and 
upright;  very  straight. 

Strigillose  (-osus).    Minutely  strigose. 

StrigoK  (-osus).  Beset  with  strigcR,  or 
sharp-pointed  and  appressed  straight 
and  stiff  hairs  or  bristles. 

Strobilaceous  (-eus),  Strobiliform  (-or- 
mis).  Relating  to  or  resembling  a 
Strobile. 

Strobile  (Strobilus).  An  inflorescence 
formed  largely  of  imbricated  scales, 
as  that  of  Hop  and  a  Fir-cone; 
303. 

Strombuliformis,  Strombuliferus.  Twist- 
ed  spirally  into  a  screw  shape,  as  the 
legumes  of  the  Screw-bean  (Proso- 
pis,  sect.  Strombocarpa)  and  of  some 
species  of  Medicago. 

Strophiole  (-iola).  An  appendage  at 
the  hilum  of  certain  seeds;  308. 

Structural  Botany,  2. 

Struma.  A  wen  or  any  cushion-like 
swelling  on  an  organ. 

Strumose  (-osus),  Strumiferous  (-us). 
Furnished  with  a  struma  or  goitre-like 
swelling. 

Stupose  (-osus).  Tow-like;  with  tufts  or 
mats  of  long  hairs. 

Style  (Stylus).  The  usually  attenuated 
portion  of  a  pistil  or  carpel  between 
the  ovary  and  stigma;  166. 

Styliform  (-ormis).    Style-shaped. 

Styliferous.     Style-bearing. 

Stylinus.    Belonging  to  the  style. 

Stylosus.  With  styles  of  remarkable 
length  or  number,  &c. 

Stylopodium.  An  enlargement  or  a  disk- 
like  expansion  at  the  base  of  a  style, 
as  in  Umbelliferse. 

Sub.  In  composition  of  Latin  words  in 
terminology,  denotes  somewhat  or 
slightly ;  as,  Subacute,  Subcordate,  that 
is  acutish,  somewhat  cordate,  &c. 

Subclass,  327. 

Subconvolute  and  Subimbricate  in  aestiva- 
tion, 137. 

Suberose  (-osus).    Of  a  corky  texture. 

Subgenus,  327. 

Submerged,  Submersed  (-us).  Growing 
under  water. 

Suborder  (Subordo),  327. 

Subpetiolar  (-arts).  Under  the  petiole, 
as  the  leaf-buds  of  PlaUnus ;  42. 

Subsection,  327. 


Subspecies.  A  group  which  is  ambigu- 
ous in  rank  between  variety  and  spe- 
cies; 320. 

Subtribe  (Subtribus),  327. 

Subulate  (-atus),  Subuliform  (-ormis). 
Awl-shaped. 

Subvariety,  327. 

Succise  (-isus).  As  if  cut  or  broken  off 
at  the  lower  end.  < 

Succubous  (-M*).  When  in  leaves 
crowded  on  a  stem  the  apex  of  each 
leaf  is  covered  by  the  base  of  the 
next  above. 

Succulent  (Succosus).    Juicy. 

Sucker.  A  shoot  of  subterranean  ori- 
gin; 53. 

Sufrutescent  (-ens).  Slightly  or  ob- 
scurely shrubby;  50. 

Suffrutex.    An  undershrub. 

Suffruticose  (-osus).  Low  and  shrubby 
at  base ;  50. 

Suffultus.     Underpropped  or  supported. 

Sukate  (-atus).     Grooved  or  furrowed. 

Super.    Above.    See  Supra. 

Superior,  Superus.  Growing  or  placed 
above ;  also  hi  a  lateral  flower  on  the 
side  next  the  axis ;  thus  the  poste- 
rior or  upper  lip  of  a  corolla  is  the 
superior;  160,  183. 

Superposed  (Superpositus).  Vertically 
over  some  other  part. 

Superposition,  179,  195. 

Supervolute  (-us),  Supervolutive  (-ivus). 
Same  as  Convolute  when  applied  to 
plaits;  139. 

Supine  (-inus).  Lying  flat  with  face  up- 
ward. 

Suppression.  Complete  abortion;  179, 
190. 

Supra.  Above;  hence  in  Latin  com- 
pounds, Supra-axillary,  above  the 
axil ;  Suprafoliaceous,  above  a  leaf,  &c. 

Supradedecompound.  Several  times  com- 
pound. 

Surculose  (-osus).    Producing  suckers. 

Surculus.  A  sucker;  a  shoot  rising  from 
a  subterranean  base ;  53. 

Sursum.  Upward;  directed  upward  or 
forward. 

Suspended  (Suspensus).  Hanging  di- 
rectly downward;  hanging  from  the 
apex  of  a  cell. 

Suspensor  of  the  embryo,  284. 

Sutural  (-alis).    Relating  to  a  suture. 

Suture  (-lira).  A  junction  or  seam  of 
union;  used  commonly  as  a  line  of 
opening;  260. 

Sword-shaped.  A  blade  with  two  sharp 
and  nearly  parallel  edges,  as  in  Iris. 


GLOSSABY. 


437 


Syconium  or  Syconut.  A  multiple  fruit 
like  that  of  the  Fig;  148,303. 

Sylvestris.    Growing  in  woods. 

Symmetrical.  Regular  as  to  number  of 
parts  or  as  to  shape.  In  the  blossom 
it  denotes  the  former;  175. 

Symmetry.  In  the  flower  relates  to 
symmetrical  disposition  of  organs  on 
the  axis;  174. 

Sympetalous  (-IM).  With  united  petals; 
same  as  Gamopetalous ;  244. 

Bymphiantherous  (-us).  Same  as  Sy- 
nantherous  and  Syngenesious. 

Symphysis.    Same  as  Coalescence. 

Symphystemonous.  With  stamens  united. 

Sympode,  Sympodium.  A  stem  made 
up  of  a  series  of  superposed  branches 
in  a  way  to  imitate  a  simple  axis ;  a 
Sympodial  stem;  55,  154. 

Synacmy.     Same  as  Synanthesis. 

Synantherous  (-us).  Stamens  coalescent 
by  their  anthers. 

Syndnthesis.  The  simultaneous  anthe- 
sis  or  readiness  of  the  anthers  and 
stigmas  of  a  blossom ;  219. 

Syncarp,  Syncarpium.  A  multiple  fruit 
such  as  a  mulberry,  or  a  fleshy  aggre- 
gate fruit,  like  that  of  Magnolia;  299. 

Syncarpous  (-us).  Composed  of  two  or 
more  united  carpels ;  261,  263. 

Syncotyledonous.  With  cotyledons  sold- 
ered together. 

Synedral.    Growing  on  the  angles. 

Synema.  The  column  of  monadelphous 
filaments,  as  in  Mallow. 

Syngenesia.  Linnsean  class  (335)  charac- 
terized by  having  the  anthers  united  or 

Syngenesious.  With  anthers  cohering 
in  a  ring;  250. 

Synonym.  A  superseded  or  unused 
name ;  354,  365. 

Synonymy.  All  that  relates  to  syno- 
nyms;  365. 

Synsepalous  (-us).  Of  coalescent  sepals ; 
same  as  Gamosepalous ;  244. 

Systematic  Botany,  2. 

Systylus.  The  coalescence  of  styles  into 
one  body. 

Tabescent  (-ens).  Wasting  or  shrivel- 
ling. 

Tail.  Any  long  and  slender  terminal 
prolongation. 

Taper-pointed.    See  Acuminate. 

Tap-root.  A  primary  descending  root 
forming  a  direct  continuation  from  the 
radicle:  31. 

Tawny.  Same  as  Fulvous;  dull  brown- 
ish-yellow. 


Taxoloffy,  Taxonomy.  Relating  to  clas- 
sification and  its  rules ;  3,  315. 

Teeth.    Any  small  marginal  lobes. 

Tegmen.    The  inner  coat  of  a  seed ;  308. 

Tela.  Latin  name  for  tissue,  cellular 
tissue,  &c. 

Teleianthus.  Same  as  perfect,  or  her- 
maphrodite-flowered. 

Tendril.  A  filiform  production  (either 
axile  or  foliar)  by  which  a  plant  may 
climb;  54. 

Tepal  (Tepalum).  A  division  of  peri- 
anth, whether  sepal  or  petal  (hardly 
ever  used). 

Teratological.  Relating  to  malforma- 
tion or  monstrous  conditions. 

Teratology.  The  science  of  monsters 
and  malformations ;  170. 

Terete  (Teres).  Round  in  the  sense  of 
having  a  circular  transverse  section. 

Tergeminate  (-atus).     Thrice  twin. 

Terminal  (-alis).  Proceeding  from  or 
belonging  to  the  end  or  apex ;  7. 

Terminology.  Same  as  Glossology;  3, 
359. 

Ternary  (-arius).  Same  as  Trimerousj 
consisting  of  three;  176. 

Ternate  ( Ternus,  Ternattu).  In  threes ; 
as  three  in  a  whorl  or  cluster. 

Tessellated  (-atus).    In  chequer-work. 

Testa.  The  outer  seed-coat,  which  it 
commonly  hard  and  brittle,  whence 
the  name,  which  answers  to  seed-shell ; 
305. 

Testaceous  (-eus).  Of  the  color  of  un- 
glazed  common  (brownish-yellow) 
pottery. 

Tetra.  In  Greek  compounds,  four; 
hence 

Tetracdrpellary  (-aril).  Of  four  car- 
pels; 261. 

Tetracdmarous  (-us),  Tetracoccut.  Of 
four  closed  carpels. 

Tetr adynamia.  Limuean  class  (335) 
which  has  the  stamens. 

Tetradynamous  (-IM).  With  four  long 
and  two  shorter  stamens ;  250. 

Tetragonal  or  Tetragonous  (-us).  Four- 
angled. 

Tetragynia.  Linnaean  artificial  order 
(337),  characterized  by  having  the 
gynoecium. 

Tetrdgynous.    Of  four  carpels  or  sty  lei. 

Tetrdmerous  (-us).  Composed  of  four 
members  in  a  circle ;  176. 

Tetrandria.    Linnsean  class  having  the 

flowers  perfect  and 

Tetrandrous.  With  four  stamens ;  249, 
334. 


438 


GLOSSARY. 


Tetrapetahui  (-*u).  With  four  petals; 
244. 

TetraphyUous  (-tw).    Four-leaved ;  243. 

Tetraquetrous  (-us).  With  four  sharp 
or  salient  angles. 

Tetrasepalous  (-us).  With  four  sepals; 
244. 

Tetrastichous  (-u*).  In  four  vertical 
ranks. 

Thalamiflorous  (-tw),  340.  With  parts 
of  the  flower  hypogynous,  or  on  the 

Thdlamus.  The  receptacle  of  a  flower ; 
167.  See  Torus. 

Thallophytes  (  Thallophyta),  341. 

Thallus.  A  stratum,  in  place  of  stem 
and  foliage. 

Theca.  A  case;  an  anther-cell  (251); 
a  spore-case,  &c.  (An  early  name  for 
the  anther,  166.) 

Tkecaphore  (-orum).  The  stipe  of  a 
carpel  (homologous  with  petiole); 
212. 

Thorn.     Same  as  spine ;  55. 

Throat.  The  orifice  of  a  gamopetalous 
corolla  or  calyx,  including  any  por- 
tion between  this  and  the  proper  tube ; 
246.  See  Faux. 

Thyrse,  Thyrsus.  A  contracted  or  ovate 
panicle;  a  mixed  inflorescence,  with 
main  axis  indeterminate,  but  the  sec- 
ondary or  ultimate  clusters  cymose; 
159. 

Tigelle,  Tigettula.  A  miniature  or  ini- 
tial stem ;  sometimes  applied  to  Cauli- 
cle  (Radicle),  sometimes  to  Plumule; 
10. 

Tinctorius.  Dyed;  used  for  dyeing; 
imparting  color. 

Tissue.    The  anatomical  fabric. 

Tomentose  (-osus).  Densely  pubescent 
with  matted  wool,  or  Tomentum. 

Tongue-shaped.  Long  and  nearly  flat, 
somewhat  fleshy,  and  rounded  at  the 
apex. 

Tooth.    See  Teeth. 

Toothed.    See  Dentate. 

Top-shaped.    Inversely  conical. 

Torose  (-osus).  Cylindrical,  with  con- 
tractions or  bulges  at  intervals. 

Tortuous  (-osus).  Bent  or  twisted  in 
different  directions. 

Torulose  (-osus).  Diminutively  or 
slightly  torose. 

Tortus.    Twisted. 

Tortilis.     Susceptible  of  twisting. 

Tontt.  The  receptacle  of  a  flower ;  167, 
211. 

Trabeculate  (-atus).     Cross-barred. 

Trachea.    A  spiral  vessel  or  duct,  named 


from  resemblance  to  the  trachea  of 
insects. 

Trachycarpous  (-us)  Rough-fruited. 

Trachyspermous.    Rough-seeded,  &c. 

Transverse  (-ersus).  Across ;  right  and 
left  as  to  bract  and  axis;  collateral; 
160. 

Trapeziform  (-ormis),  Trapezoid.  Un- 
symmetrically  four-sided,  like  a  tra- 
pezium. 

Tree.  A  woody  plant  with  an  elevated 
trunk. 

Tri-.  In  compound  words,  both  Lathi 
and  Greek,  denotes  three  or  triple. 

Triachanium.  A  fruit  like  a  cremocarp 
but  of  three  carpels. 

Triadelphous,  Triadelphia.  With  fila- 
ments in  three  sets ;  250. 

Triandria.  Linnaean  class  (334)  with  the 
flowers. 

Tridndrous.    With  three  stamens;  249. 

Triangular  (-aris),  Triangulatus.  Three- 
angled. 

Tridnthous  (-us).    Three-flowered. 

Tribe.  Group  superior  to  genus,  In- 
ferior to  order ;  326. 

TricarpeUary  (-arts).  Of  three  carpels ; 
261. 

Tricarpous  (-tw).  Consisting  of  three 
fruits  or  carpels. 

Tricephalous  (-us).    Bearing  three  heads. 

Trichocarpous  (-tw).    Hairy-fruited. 

Trichodes.    Resembling  hair. 

Trichdtomous  (-tw).  Three-forked; 
branched  into  three  divisions. 

Trichome  (Trichdma).  Any  outgrowth 
of  the  epidermis,  such  as  a  hair  or 
bristle;  209. 

Tricticcous  (-tw).  Consisting  of  three 
cocci. 

Tricolor.    Three-colored. 

Tricuspidate  (-atus).  Tipped  with  three 
cusps  or  pointed  tips. 

Tridentate  (-atus).    Three-toothed. 

Tridigitate.    Thrice  digitate. 

Triduus.    Lasting  for  three  days. 

Triennial  ( Triennis).  Lasting  for  three 
years. 

Trifarious  (-ius).  Facing  three  ways, 
in  three  vertical  ranks. 

Trifid  (Trifidus).    Three-cleft. 

Trifoliate  (-ius,  Trifoliatus).  Three- 
leaved. 

Trifoliolate  (-atus).    Of  three  leaflets. 

Trifurcate  (-atus).  Divided  into  three 
forks  or  branches. 

Trigamous  (-us).  Bearing  three  kind* 
of  flowers. 

Trigonous  (-ut),  Trigonal.  Three-«ngled. 


GLOSSARY. 


439 


Trigyma.  Linnsean  artificial  order  with 
Trigynout,  »'.  «.  three-styled  flowers ; 
337. 

Trihilatus.  Having  three  apertures,  as 
in  some  grains  of  pollen. 

Trijugate  ( Trijugus).  With  three  pairs 
of  leaflets  or  pinnae. 

Trilobate  (Trilobus).     Three-lobed. 

Trilocular  (-aril).     Three-celled. 

Trimerous  (-MS).  Three-membered  parts 
in  threes;  176. 

Trimestris.  Lasting  for  or  maturing  in 
three  months. 

Trimorphous,  Trimorpism.  Occurring 
under  three  forms;  236. 

Trinervate  ( Trinervius).    Three-nerved. 

Trinodal.    Of  three  nodes  or  joints. 

Triaecia.  Linnsean  artificial  order  with 
the  flowers. 

Tricecious  or  Trioicous  (-us).  Having 
staminate,  pistillate  and  perfect  flowers 
(or  three  kinds  of  flowers  as  to  sex), 
334;  on  three  distinct  plants. 

Triovulate  (-atus).   Having  three  ovules. 

Tripartite  (-ibilis).  Tending  to  split 
into  three  portions. 

Tripartite  (-itus).    Three-parted. 

Tripetaloid  (-oideus).  As  if  three-pet- 
alled. 

Tripetalous  (-MS).    Having  three  petals. 

Triphyllous  (-MS).     Three-leaved ;  243. 

Tripinnate  (-atus).   Thrice  pinnate ;  104. 

Tripinnatifid  (-idus).     Thrice  pinnatifid. 

Triple-ribbed  or  nerved.  With  midrib 
dividing  into  three,  or  sending  off  on 
each  side  a  strong  branch,  above  the 
base  of  the  blade;  93. 

Triplinerved  (Triplinervius).  Same  as 
Triple-nerved,  Triple-ribbed;  93. 

Tripterous  (-us).     Three-winged. 

Triquetrous  (Triqueter).  Three-edged; 
with  three  salient  angles. 

Triquinate  (-atus).  Divided  first  into 
three  then  into  five. 

Trisected  (-us).  Divided  into  three  por- 
tions; 99. 

Trisepalous  (-MS).     Of  three  sepals. 

Triserial  (-alis),  Triseriate  (-atus).  In 
three  horizontal  ranks  or  series. 

Tristdchyus.    Three-spiked. 

Tristichous  (-us).  In  three  vertical 
ranks;  122. 

Tristigmatic.    With  three  stigmas. 

Tristis.     Dull  colored. 

Tristylous  (-us).    Having  three  styles. 

Trifulcate  (-atus).     Three-grooved. 

Triternate  (atus).     Thrice  ternate;  104. 

Trivial  names,  Nomina  trivialia.  Com- 
mon or  vulgar  names ;  used  by  Lin- 


naeus for  specific  names  of  a  single 
word ;  346,  362. 

Trochlear  (-earis).     Pulley-shaped. 

Trophosperm  (Trophospermium).  Name 
for  the  Placenta;  261. 

Trumpet-shaped.  Tubular,  with  a  dilat- 
ed orifice. 

Truncate  (-atus).  As  if  cut  off  at  the 
end;  97. 

Trunk  ( Truncus).    A  main  stem. 

Tryma.  A  drupaceous  nut,  with  exo- 
carp  at  length  dehiscent  or  otherwise 
separating,  such  as  walnut  and  hick- 
ory nut. 

Tubceformis.    Trumpet-shaped. 

Tube  (  Tubus).  Any  hollow  elongated 
body  or  part  of  an  organ ;  245. 

Tuber.  A  thickened  and  short  subter- 
ranean branch,  beset  with  buds  or 
eyes;  59. 

Tubercle  ( Tuberculum).  A  small  tuber 
or  an  excrescence:  or  something  be- 
tween a  tuber  and  a  root ;  60. 

Tuberculate  (-atus).  Beset  with  knobby 
projections  or  excrescences. 

Tuberiferous.    Bearing  tubers. 

Tubular,  Tubulosus  (-ose).  Having  a 
tube;  tube-shaped;  248. 

Tubuliflorus  (-us).  When  the  flowers  of 
a  head  have  only  tubular  corollas. 

Tunicate  (-MS).    Having  coats  (tunics). 

Turbinate  (-atus).        Top-shaped. 

Turion,  (Turio).  A  scaly  sucker  or 
shoot  from  the  ground ;  41. 

Turnip-shaped.    See  Napiform. 

Twin.  In  pairs.  See  Geminate,  Didy- 
mous. 

Tinning.  Winding  spirally  and  so 
climbing  ( Twiners);  51. 

Twisted.    Contorted. 

Two-lipped.    See  Bilabiate. 

Type.    The  ideal  plan  or  pattern. 

Typical.  Representing  the  plan  or 
type. 


Uliginose  (-osus).    Growing  In  swamps. 

Ulndris.  Of  the  length  of  the  ulna  or 
fore-arm. 

Umbel  ( Umbella).  An  inflorescence 
(properly  of  the  indeterminate  type) 
in  which  a  cluster  of  pedicels  spring 
all  from  the  same  point,  like  rays  of 
an  umbrella;  146. 

Umbellate  (-atus),  UmbeUiform  (-ormis). 
In  or  like  umbels. 

Umbdlet.  A  partial  or  secondary  um- 
bel; 150. 

Umbelliferous  (-us).    Bearing  umbela- 


440 


GLOSSARY. 


Umbettula.     A  partial  or  secondary  um- 
bel, or  umbellet;  150. 
Umbilicate  (-atus).     Depressed  in  the 

centre,  navel  like. 
Umbilicus.    The  hilum  of  a  seed. 
Umbonate  (-atus).     Bearing  an    Umbo 

or  loss  in  the  centre. 
Umbrdculiform   (-ormis).     Having  the 

general  form  of  an  umbrella. 
Umbrosus.    Growing  in  shady  places. 
Unarmed.    Destitute  of  prickles,  spines, 

or  other  armature. 

Uncate  (-atus),  Uncinate,  (-atus),  Unci^- 

form   (-ormis).       Hooked;    bent    or 

curved  at  tip  in  the  form  of  a  hook. 

Unddlis.     An  inch  (uncia)  in  length. 

Undate     (-atus)    or    Undulate    (-atus). 

Wavy;  98. 

Undenhrub.    A  very  low  shrub ;  50. 
Unequally   pinnate.      See    Impari-pin- 

nate. 
Unguiculate    (-atus).      Contracted    at 

base  into  an 
Unguis.    A  claw,  or  stalk-like  base  of  a 

petal,  &c. ;  245. 

Uni-.    In  Latin  compound,  one ;  as 
Unicellular.    Of  one  cell;   Unicolor,  of 

one  color,  &c. 

Unicus.    Singly  or  single,  solitary. 
Uniflorous  (-us).    One-flowered. 
Unifoliate  (-atus).    One-leaved. 
Unifoliolate,  of  one  leaflet ;  102. 
Unijugate    (Unijugus).      Of  one  pair; 

102. 

Unilabiate  (-atus).    One-lipped,  like  the 
corolla    of  Acanthus,   in  which    the 
upper  lip  is  obsolete. 
Unilateral    (-alis).     One-sided;    either 
originating    on    or    more    commonly 
turned  all  to  one  side  of  an  axis. 
Unilocular  (-aris).     One-celled. 
Uninervate     ( Uninervis,     Uninervius). 

One-nerred. 

Uniovulate  (-atus).    Having  only  a  soli- 
tary ovule. 
Uniparous.    Bearing  one ;  as  a  cyme  of 

one  axis  or  branch ;  152,  155. 
Uniserial    (-ialis),    Uniseriate    (-atus). 

In  one  horizontal  row  or  series. 
Unisexual    (-alis,    Unisexus).      Of   one 
sex;  having  stamens  only  or  pistils 
;only;  191. 
Univalved  (  Univalvis).    Of  one  piece  or 

valve. 

Urceolate    (-atus).      Hollow   and    con- 
tracted at  or  below  the  mouth,  like  an 
urn  or  pitcher  (  Urceolus). 
Urens.    Stinging,  in  the  manner  of  net- 
tles. 


Utiicle  (  Utriculus).  A  small  bladdery 
pericarp ;  295.  Or  any  small  bladder- 
shaped  body  or  appendage;  also  a 
synonym  of  a  cell  of  parenchyma. 

Utricular  (-arts),  Utriculate  (-atus), 
Utriculiform  (-ormis),  Utriculose 
(-osus).  Having  or  consisting  of 
utricles,  or  bladder-like  in  appear- 


Vacillans.  Swinging  free,  as  the  anth- 
ers of  Grasses  on  their  filaments. 

Vdcuus.  Void  or  empty  of  the  proper 
contents. 

Vagina.    A  sheath,  as  of  a  leaf,  &c. 

Vaginate.     Sheathed. 

Vatteculce.  The  intervals  or  grooves 
between  the  ridges  or  ribs  of  the  fruit 
Umbelliferse. 

Valvate  (-atus),  Valvular  (-aris). 
Opening  as  if  by  doors  or  valves,  as 
do  most  dehiscent  fruits  (capsules), 
and  some  anthers;  also  the  parts  of 
a  flower-bud  when  they  exactly  meet 
without  overlapping ;  135. 

Valve  (  Valva).  One  of  the  pieces  into 
which  a  capsule  splits,  288. 

Valved.  Same  as  valvate:  hence  3 
-valved,  5-valved,  many  valved,  &c. 

Vdlvula.  A  diminutive  valve.  Also 
used  (after  Linnaeus)  for  the  inner  or 
flower-glumes  of  Grasses. 

Variegated  (-atus).  Irregularly  colored ; 
in  patches  of  color. 

Variety  (  Varietas).  A  sort  or  modifi- 
cation subordinate  to  species ;  318. 

Variolate,  Varioldris.  Marked  as  if 
by  the  pustules  or  pittings  of  small- 
pox. 

Vascular  (-aris).  Relating  to  or  fur- 
nished with  vessels  ( Vasa)  or  ducts. 

Vascular  Plants  (  Vasculares),  340. 

Vdsculum.  Same  as  Ascidium.  Also 
the  botanists1  collecting  box ;  372. 

Vasiform  (-ormis).  In  the  form  of  a 
vessel,  duct,  &c. 

Veined.  Furnished  or  traversed  with 
nbro-vascular  bundles  or  threads,  es- 
pecially with  those  which  divide  and 
are  reticulated. 

Veins  (  Vena).  In  general  any  ramifi- 
cations or  threads  of  fibro-vascular 
tissue  in  a  leaf  or  any  flat  organ; 
especially  (as  distinguished  from 
nerves )  those  which  divide  or  branch ; 
92. 

Veinless.    Destitute  of  veins. 

Veinlet  (  Venula).     One  of  the  ultimata 


GLOSSARY. 


441 


or  smaller  ramifications  of  a  vein  or 
rib;  93. 

V elate  (-atut).     Veiled. 

Velutinous  (Velutinus).  Velvety:  the 
surface  covered  with  a  soft  coating  of 
fine  and  close  silky  pubescence,  or 
relumen. 

Venation  ( Venatio).  The  mode  of  vein- 
ing;  90. 

Venenatttt,  Ventnonu.    Poisonous. 

Venose  (-osus)  Veiny;  abounding  in 
veins  or  network. 

Ventral  (-alis).  Belonging  to  the  an- 
terior or  inner  face  of  a  carpel,  &c. ; 
the  opposite  of  dorsal. 

Ventricose  (-osus).  Swelling  unequally 
or  inflated  on  one  side. 

Ventriculose  (-ona).  Minutely  ventri- 
cose. 

Venulote  (-otut).  Abounding  with  vein- 
lets  or  venuUe. 

Vermicular  (-arit).     Worm-shaped. 

Vernal  ( Verndlit).  Appearing  hi  spring. 

Vernation  (-atio).  The  disposition  of 
parts  in  a  leaf-bud ;  132. 

Vemicose  (-osus).    As  if  varnished. 

Verrucose  (-osus).  Covered  with  warts 
(verruca)  or  wart-like  elevations. 

Vertatile  ( Versatile).  Swinging  to  and 
fro;  turning  freely  on  its  support; 
_^  253. 

Versicolor.  Changing  color,  or  of  more 
than  one  tint  or  color. 

Vertex.    The  apex  of  an  organ. 

Vertical  (-alit).  Perpendicular  to  the 
horizon ;  longitudinal. 

Verticil  (-illus).    A  whorl;  6. 

Verticittaster.  A  false  whorl,  composed 
of  a  pair  of  opposite  cymes ;  159. 

VerticMastrate.  Bearing  or  arranged 
in  Verticillasters. 

Verticillate  (-atut,  -arts).  Disposed  in  a 
whorl ;  6,  119,  120. 

Vescicle  (-icula).  A  small  bladder  or 
air-cavity. 

Vesicular  (-arit),  Vesiculote  (-otut).  As 
if  composed  of  little  bladders. 

Vespertine  (  Vespertinus).  Appearing  or 
expanding  in  early  evening. 

Vettels  ( Vatas).    See  Ducts. 

Vexillary  (-arit),  Vexillar,  137.  Per- 
taining to  the 

Vexillwn.  The  standard  or  large  pos- 
terior petal  of  a  papilionaceous  corolla; 
184. 

ViUose  (otut)  or  ViUout.  Bearing  shaggy 
or  long  and  soft  (not  interwoven)  hairs 
or  Villi. 

Vimineout(-eut).  Bearing  long  and  flex- 


ible twigi,  like  thon  used  for  wicker 

work. 
Vine.    Any  trailing  or  climbing  stem : 

originally  that    of    the    Grape    from 

which  wine  is  made. 
Vinedlit.    Growing  in  vineyards. 
Violaceous  (-eut).    Violet-colored. 
Virent.    Green,  or  evergreen. 
Viresceni.     Greenish  or  turning  green. 
Virgate  (-atus).    Wand-shaped,  or  like 

a  rod ;  slender,  straight,  and  erect. 
Virgultum.     A  vigorous  twig  or  shoot. 
Viridetcent  (-ent).    Same  as  Virescens. 
Viridis.    Green. 
Viridultu.   Greenish. 
Vinous.    Venomous. 
Viscid  (-idus),   Vitcous  (-otut).    Sticky 

from  a  tenacious  coating  or  secretion. 
Vitellinus.    The  yellow  hue  of  the  yolk 


Vitellut.  Name  formerly  given  to  the 
peculiar  albumen  which  is  in  some 
cases  deposited  within  the  embryo-sac. 

Viticulose  (-osus).  Sarmentaceous ;  pro- 
ducing vine-like  twigs  or  suckers, 
viticula. 

Vittte.  The  fillets  or  stripes  (oil-tubes) 
of  the  pericarp  of  most  Umbelliferae, 
which  contain  an  aromatic  or  peculiar 
secretion. 

Vitiate  (-atus).  Bearing  vitte ;  or  with 
any  longitudinal  stripes. 

Viviparous  ( -us ) .  Germinating  or  sprout- 
ing from  seed  or  bud  while  on  the 
parent  plant. 

Voluble  (Volubilis).  Twining  round  a 
support ;  51. 

Volutus.    Rolled  up  in  any  way. 

Volva.  A  wrapper  or  external  covering, 
especially  that  of  many  Fungi. 


Wavy.    See  Undulate. 

Waxy.  Resembling  beeswax  in  consist- 
ence or  appearance. 

Wedge-shaped  or  Wedge-form.  See 
Cuneate;  95. 

Wheel-shaped.     See  Rotate. 

Whorl.  Arranged  in  a  circle  round  an 
axis;  a  Verticil;  6. 

Whorled.    Disposed  in  whorls. 

Wild.  Growing  without  cultivation,- 
spontaneous. 

Winy.  See  Ala.  Any  Membraneous 
or  thin  expansion  by  which  an  organ 
is  bordered,  surrounded,  or  otherwise 
augmented.  Also  the  two  lateral 
petals  of  a  papilionaceous  corolla  are 
termed  wings ;  185. 


442 


GLOSSARY. 


Winged.     See  Alate :  bearing  a  wing  or 

wings. 

Withering.    See  Marcescent. 
Wood.    The  hard  part  of  a  stem,  &c., 

mainly  composed  of 
Wood-cell*,  Woody  fibre  or  tissue,  68. 
Woolly.     See  Lanate  and   Tomentose: 

clothed    with    long  and  tortuous  or 

matted  hairs. 

Xanthos.  Greek  for  yellow  in  com- 
pounds, such  as  Xanthvphyll,  the  yel- 
low coloring  matter  in  leaves. 


Xenogamy.  Fecundation  of  the  ovules  of 
a  flower  by  pollen  from  some  other 
plant  of  the  same  species;  cross-fer- 
tilization; 216. 

Xylinns.    Woody,  pertaining  to  wood. 


Zoospore.  One  of  the  free-moving  spores 

of  the  lower  Cryptogams. 
Zygomorphous  (-us).    That  which  can  be 

bisected  in  only  one  plane  into  similar 

halves;  175. 


ADDENDA. 

Antidromous,  Antidromy.    When  the  course  of  a  spiral  is  reversed,  157. 
Infertile  (-Hit).     Said  of  a  pistil  or  flower  which  fails  to  set  fruit. 
Polyembryony.    The  production  of  two  or  more  embryos  in  a  seed.  284. 
Saprophytes  (-yta).    Plants  feeding  upon  decaying  vegetable  OT  animal  matter 


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